Hubbry Logo
Morus (plant)Morus (plant)Main
Open search
Morus (plant)
Community hub
Morus (plant)
logo
8 pages, 0 posts
0 subscribers
Be the first to start a discussion here.
Be the first to start a discussion here.
Morus (plant)
Morus (plant)
Morus (plant)
View on Wikipedia
from Wikipedia
"Mulberry" redirects here. For other plants called mulberry, see List of plants known as mulberry. For other uses, see Mulberry (disambiguation).
For the bird genus, see Gannet.

Mulberry
Morus nigra
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Rosales
Family: Moraceae
Tribe: Moreae
Genus: Morus
L.
Species

See text.

Morus, a genus of flowering plants in the family Moraceae, consists of 19 species of deciduous trees commonly known as mulberries, growing wild and under cultivation in many temperate world regions.[1][2][3][4] Generally, the genus has 64 subordinate taxa,[5] though the three most common are referred to as white, red, and black, originating from the color of their dormant buds and not necessarily the fruit color (Morus alba, M. rubra, and M. nigra, respectively), with numerous cultivars and some taxa currently unchecked and awaiting taxonomic scrutiny.[6][5] M. alba is native to South Asia, but is widely distributed across Europe, Southern Africa, South America, and North America.[2] M. alba is also the species most preferred by the silkworm. It is regarded as an invasive species in Brazil, the United States and some states of Australia.[2][7]

The closely related genus Broussonetia is also commonly known as mulberry, notably the paper mulberry (Broussonetia papyrifera).[8]

Despite their similar appearance, mulberries are not closely related to raspberries or blackberries. All three species belong to the Rosales order. But while the mulberry is a tree belonging to the Moraceae family (also including the fig, jackfruit, and other fruits), raspberries and blackberries are brambles and belong to the Rosaceae family.[9]

Description

[edit]

Mulberries are fast-growing when young, and can grow to 24 metres (79 feet) tall.[2][6] The leaves are alternately arranged, simple, and often lobed and serrated on the margin. Lobes are more common on juvenile shoots than on mature trees.[2][6] The trees can be monoecious or dioecious.[6]

The mulberry fruit is a multiple, about 2–3 centimetres (3⁄4–1+1⁄4 inches) long.[2][6] Immature fruits are white, green, or pale yellow.[6] The fruit turns from pink to red while ripening, then dark purple or black, and has a sweet flavor when fully ripe.[2][6]

  • Clusters (inflorescences) of unopened male flower buds
    Clusters (inflorescences) of unopened male flower buds
  • Female catkins
    Female catkins
  • Young mulberry fruit clusters
    Young mulberry fruit clusters
  • Immature fruit
    Immature fruit
  • Unripe white mulberries
    Unripe white mulberries
  • Berries on branches in Eastern Oklahoma
    Berries on branches in Eastern Oklahoma
  • Mulberry in southern Brazil
    Mulberry in southern Brazil
  • Long mulberry
    Long mulberry
  • Semi-ripe mulberries on a mulberry leaf
    Semi-ripe mulberries on a mulberry leaf
  • Autumn foliage
    Autumn foliage

Taxonomy

[edit]

The taxonomy of Morus is complex and disputed. Fossils of Morus appear in the Pliocene record of the Netherlands.[10] Over 150 species names have been published, and although differing sources may cite different selections of accepted names, fewer than 20 are accepted by the vast majority of botanical authorities. Morus classification is further complicated by widespread hybridisation, wherein the hybrids are fertile.[citation needed]

The following species are accepted:[11]

Distribution

[edit]
Mulberry fruit in Libya

Black, red, and white mulberries are widespread in Southern Europe, the Middle East, Central Asia, Northern Africa, and the Indian subcontinent, where the tree and the fruit have names under regional dialects.

Black mulberry was imported to Britain in the 17th century in the hopes that it would be useful in the cultivation of silkworms.[12] It was much used in folk medicine, especially in the treatment of tapeworms.[13]

The United States has native red mulberries,[14] as well as imported black and white mulberries. In North America, the white mulberry is considered an invasive exotic and has taken over extensive tracts from native plant species, including the red mulberry.[2][15]

Mulberries are also widespread in Greece, particularly in the Peloponnese, which in the Middle Ages was known as Morea, deriving from the Greek word for the tree (μουριά, mouria).

Australia has two types of native mulberries: Hedycarya angustifolia, and Pipturus argenteus,[16] which are both from different families to Moraceae. The exotic black, red and white Morus mulberries are also commonly grown in Australian backyards.[17] White mulberry is considered an environmental weed in the states of Queensland and New South Wales.[7]

Cultivation

[edit]
A mulberry tree in England

Mulberries can be grown from seed, and this is often advised, as seedling-grown trees are generally of better shape and health.[citation needed] Mulberry trees grown from seed can take up to ten years to bear fruit. Mulberries are most often planted from large cuttings, which root readily. The mulberry plants allowed to grow tall have a crown height of 1.5 to 1.8 m (5 to 6 ft) [citation needed] from ground level and a stem girth of 10–13 cm (4–5 in). They are specially raised with the help of well-grown saplings 8–10 months old of any of the varieties recommended for rainfed areas like S-13 (for red loamy soil) or S-34 (black cotton soil), which are tolerant to drought or soil-moisture stress conditions. Usually, the plantation is raised and in block formation with a spacing of 1.8 by 1.8 m (6 by 6 ft), or 2.4 by 2.4 m (8 by 8 ft), as plant-to-plant and row-to-row distances. The plants are usually pruned once a year during the monsoon season to a height of 1.5–1.8 m (5–6 ft) and allowed to grow with a maximum of 8–10 shoots at the crown.[citation needed]

Mulberry tree scion wood can easily be grafted onto other mulberry trees during the winter, when the tree is dormant. One common scenario is converting a problematic male mulberry tree to an allergy-free female tree, by grafting all-female mulberry tree scions to a male mulberry that has been pruned back to the trunk.[18] However, any new growth from below the graft(s) must be removed, as they would be from the original male mulberry tree.[19]

Toxicity and allergenicity

[edit]

All parts of the plant besides the ripe fruit can exude a milky sap (latex) which is mildly toxic if ingested, causing digestive distress and, at larger doses, hallucinations.[20][21][22] It is also an irritant and may cause a skin rash on contact.[23] Unripe green fruit may cause nausea, cramps, and be hallucinogenic.[24] The berries have a laxative effect; too many will cause diarrhea.[25]

Some North American cities have banned the planting of mulberries because of the large amounts of pollen they produce, posing a potential health hazard for some pollen allergy sufferers.[26] Only the male mulberry trees produce pollen; this lightweight pollen can be inhaled deeply into the lungs, sometimes triggering asthma.[27][28] Conversely, female mulberry trees produce all-female flowers, which draw pollen and dust from the air. Because of this pollen-absorbing feature, all-female mulberry trees have an OPALS allergy scale rating of just 1 (lowest level of allergy potential), and some consider it "allergy-free".[27]

Nutrition

[edit]
Raw mulberries
Nutritional value per 100 g (3.5 oz)
Energy180 kJ (43 kcal)
9.8
Sugars8.1
Dietary fiber1.7
0.39
1.44
Vitamins and minerals
VitaminsQuantity
%DV†
Vitamin A equiv.
0%
1 μg
Thiamine (B1)
2%
0.029 mg
Riboflavin (B2)
8%
0.101 mg
Niacin (B3)
4%
0.62 mg
Vitamin B6
3%
0.05 mg
Folate (B9)
2%
6 μg
Vitamin C
40%
36.4 mg
Vitamin E
6%
0.87 mg
Vitamin K
7%
7.8 μg
MineralsQuantity
%DV†
Calcium
3%
39 mg
Iron
10%
1.85 mg
Magnesium
4%
18 mg
Phosphorus
3%
38 mg
Potassium
6%
194 mg
Sodium
0%
10 mg
Zinc
1%
0.12 mg
Other constituentsQuantity
Water87.68 g
US Department of Agriculture FoodData Central entry[29]
†Percentages estimated using US recommendations for adults,[30] except for potassium, which is estimated based on expert recommendation from the National Academies.[31]

Raw mulberries are 88% water, 10% carbohydrates, 1% protein, and less than 1% fat. In a 100-gram (3.5-ounce) reference amount, raw mulberries provide 43 calories, 44% of the Daily Value (DV) for vitamin C, and 14% of the DV for iron; other micronutrients are insignificant in quantity.[29]

Uses

[edit]

Agricultural

[edit]

The leaves are harvested three or four times a year by a leaf-picking method under rain-fed or semi-arid conditions, depending on the monsoon. The leaves are useful as animal fodder. The tree branches pruned in the fall (autumn; after the leaves have fallen) are cut and used to make durable baskets supporting agriculture and animal husbandry.[citation needed]

Culinary

[edit]

As the fruit matures, mulberries change in texture and color, becoming succulent, plump, and juicy, resembling a blackberry.[6] The color of the fruit does not distinguish the mulberry species, as mulberries may be white, lavender or black in color. The fruit of the black mulberry (native to southwest Asia) and the red mulberry (native to eastern North America) have distinct flavors.[32] White mulberry fruits are typically sweet, but not tart, while red mulberries are usually deep red, sweet, and juicy. Black mulberries are large and juicy, with balanced sweetness and tartness.[6] The fruit of the East Asian white mulberry – a species extensively naturalized in urban regions of eastern North America – has a different flavor, sometimes characterized as refreshing and a little tart, with a bit of gumminess to it and a hint of vanilla.[32][better source needed]

Mulberries are used in pies, tarts, wines, cordials, and herbal teas.[2][6] Jams and sherbets are often made from the fruit in the Old World. In spring, new tender twigs are semisweet and can be eaten raw or cooked.[33]

In Armenia, mulberries are common, with wide usage in homemade oghi (moonshine) production, known as tti oghi, and syrup known as doshab.[34]

Supplement

[edit]

The fruit and leaves are sold in various forms as dietary supplements.[23][35]

Silk industry

[edit]
A silkworm, Bombyx mori, feeding on a mulberry tree

Mulberry leaves, particularly those of the white mulberry, are ecologically important as the sole food source of the silkworm (Bombyx mori, named after the mulberry genus Morus), the cocoon of which is used to make silk.[36][37] The wild silk moth also eats mulberry.[38][39] Other Lepidoptera larvae—which include the common emerald, lime hawk-moth, sycamore moth, and fall webworm—also eat the plant.[40]

The Ancient Greeks and Romans cultivated the mulberry for silkworms; at least as early as 220 AD, Emperor Elagabalus wore a silk robe.[41] English clergy wore silk vestments from about 1500 onwards.[41] Mulberry and the silk industry played a role in colonial Virginia.[41]

Pigment

[edit]

Mulberry fruit color derives from anthocyanins, which have unknown effects in humans.[42] Anthocyanins are responsible for the attractive colors of fresh plant foods, including orange, red, purple, black, and blue.[42] These colors are water-soluble and easily extractable, yielding natural food colorants.[2] Due to a growing demand for natural food colorants, they have numerous applications in the food industry.[3][42]

A cheap and industrially feasible method has been developed to extract anthocyanins from mulberry fruit that could be used as a fabric dye or food colorant of high color value.[2] Scientists found that, of 31 Chinese mulberry cultivars tested, the total anthocyanin yield varied from 148 to 2725 mg/L of fruit juice.[43] Sugars, acids, and vitamins of the fruit remained intact in the residual juice after removal of the anthocyanins, indicating that the juice may be used for other food products.[43][2]

Mulberry germplasm resources may be used for:[3][2][44]

  • exploration and collection of fruit yielding mulberry species
  • their characterization, cataloging, and evaluation for anthocyanin content by using traditional, as well as modern, means and biotechnology tools
  • developing an information system about these cultivars and varieties
  • training and global coordination of genetic stocks
  • evolving suitable breeding strategies to improve the anthocyanin content in potential breeds by collaboration with various research stations in the field of sericulture, plant genetics, and breeding, biotechnology and pharmacology

Paper

[edit]

During the Angkorian age of the Khmer Empire of Southeast Asia, monks at Buddhist temples made paper from the bark of mulberry trees. The paper was used to make books, known as kraing.[45]

Tengujo is the thinnest paper in the world. It is produced in Japan and made with kozo (stems of mulberry trees).[46] Traditional Japanese washi paper is often created from parts of the mulberry tree.[47]

Wood

[edit]

The wood of mulberry trees is used for barrel aging of Țuică, a traditional Romanian plum brandy.[48]

Culture

[edit]
Mulberry Tree by Vincent van Gogh

A Babylonian etiological myth, which Ovid incorporated into his Metamorphoses, attributes the reddish-purple colour of the mulberry fruits to the tragic deaths of the lovers Pyramus and Thisbe. Meeting under a mulberry tree (probably the native Morus nigra),[49] Thisbe dies by suicide by sword after Pyramus does the same, he having believed, on finding her bloodstained cloak, that she was killed by a lion. Their splashed blood stained the previously white fruit, and the gods forever changed the mulberry's colour to honour their forbidden love.[49]

In the Old Testament's 1 Maccabees, the Seleucids used the "blood of grapes and mulberries" to provoke their war elephants in preparation for battle against Jewish rebels.[50][51] In the New Testament, Luke 17:6 refers to the power of a little faith to move a mulberry tree into the sea.[52]

The nursery rhyme "Here We Go Round the Mulberry Bush" uses the tree in the refrain, as do some contemporary American versions of the nursery rhyme "Pop Goes the Weasel".[53][54]

Vincent van Gogh featured the mulberry tree in some of his paintings, notably The Mulberry Tree (Mûrier, 1889, now in Pasadena's Norton Simon Museum). He painted it after a stay at an asylum, and he considered it a technical success.[55]

References

[edit]
[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Morus (plant)

View on Grokipedia
from Grokipedia
Morus is a of flowering in the family , comprising 17 accepted of trees and shrubs known collectively as mulberries. These are characterized by their milky sap, alternate simple leaves that vary from entire to deeply lobed with toothed margins, and distinctive aggregate fruits formed from fused drupes, which ripen to colors ranging from white and pink to red and black depending on the . Native primarily to the temperate and subtropical regions of the , the exhibits a disjunct distribution with approximately 13 in (particularly East and ) and three in the , while one , M. nigra, is endemic to southwestern . Taxonomically, Morus belongs to the tribe Moreae within Moraceae and was first described by Carl Linnaeus in 1753, though its systematics have been challenging due to high morphological variability, hybridization, and incomplete lineage sorting. Phylogenetic studies indicate that the genus is monophyletic in its strict sense (Morus s.s.), excluding genera like Afromorus and Paratrophis, and originated around the Eocene–Oligocene boundary approximately 34 million years ago, with diversification accelerating in the early Miocene linked to the development of temperate forests. Key species include Morus alba (white mulberry), widely cultivated for its glossy leaves and role in sericulture; M. nigra (black mulberry), prized for its flavorful dark fruits and ornamental value; and M. rubra (red mulberry), a North American native valued for its edible berries. Economically, the genus is most notable for its importance in , as the leaves of M. alba serve as the primary source for the silkworm , supporting about 90% of global raw production and providing livelihoods in rural economies across . Mulberry fruits are also consumed fresh, dried, or processed into juices, wines, and dyes, while various plant parts exhibit medicinal properties, including , antidiabetic, and effects attributed to bioactive compounds like and . Additionally, mulberries contribute to environmental benefits such as and wildlife habitat, though some species like M. alba have become invasive in non-native regions.

Morphology and Description

Physical Characteristics

Morus species are deciduous trees or shrubs that typically reach heights of 10 to 20 meters, though some can grow up to 25 meters or more under optimal conditions. They exhibit a dense, spreading crown that is often wider than the tree's height, with a short bole and low-branching habit in cultivation. These plants contain milky sap in their stems and leaves, a characteristic trait of the family. Most species are dioecious, with male and female flowers on separate plants, though occasional monoecious individuals occur, such as in M. laevigata. The leaves of Morus are alternate, simple to palmately lobed, and measure 5 to 20 cm in length, with serrated or toothed margins. They feature 3 to 5 primary veins arising from the base and pinnate secondary venation, showing variability in shape even within the same —often unlobed in M. alba and more frequently lobed in M. rubra. Leaf surfaces vary by : glossy and smooth in M. alba, pubescent on the lower surface in M. rubra, and matte with a scabrous texture in M. nigra. Stems are initially smooth with gray bark in young , becoming rough and deeply fissured with age on mature trunks. Branchlets range from glabrous to densely pubescent depending on the , and winter buds consist of 3 to 6 imbricate scales. Flowers are inconspicuous and arranged in axillary catkins or spikes, blooming in spring; male flowers form elongate spikes, while female flowers are in shorter spikes or capitate clusters, with dioecious arrangements predominant. The fruits develop as aggregate fruits composed of multiple drupes, typically 1 to 3 cm long, with colors varying by —white to pinkish or purplish in M. alba, dark red to purplish in M. rubra, and black in M. nigra.

Growth and Life Cycle

Morus species exhibit rapid juvenile growth, typically achieving heights of 3 to 4 meters within the first 4 to 6 years, equating to an average rate of 0.5 to 1 meter per year, before slowing in maturity. In the wild, these trees generally live 30 to 70 years, though cultivated specimens can exceed 100 years, as evidenced by historical examples such as a tree at dating to 1785. In species such as M. rubra, optimum seed production occurs between 30 and 85 years of age in forest stands. The life cycle of Morus begins with from , which can occur under favorable moist conditions, followed by a phase of vegetative growth lasting 2 to 3 years characterized by rapid shoot and extension. Reproductive maturity is reached at 3 to 5 years, when trees begin producing flowers and fruits, with open-grown individuals potentially fruiting as early as 2 years. Vegetative propagation via sprouting after disturbance allows for perennial persistence, enabling recovery from environmental stresses. Seasonally, Morus species flush new leaves in spring following winter , marked by complete drop in temperate zones. Fruiting occurs in summer, from May to in temperate regions, aligning with peak activity and environmental warmth. This cycle reflects adaptation to habits, with providing resilience to cold and aiding in resource conservation. Growth is optimized in full sun, where seedlings show up to 53% higher rates compared to shaded conditions, though partial shade is tolerated with reduced vigor. The root system features a deep penetrating 2 to 4 meters and extensive lateral spread of 7 to 13 meters, facilitating and . In dioecious Morus species, branching patterns form stout, spreading crowns, with monoecious variants occasionally exhibiting mixed-sex catkins on the same branches; however, sex-specific growth rate differences remain undocumented in primary literature. Annual wood production contributes to durable heartwood used historically in implements, though quantitative yields vary by and site.

Taxonomy and Phylogeny

Classification History

The genus Morus was established by in his in 1753, initially encompassing species such as M. alba, M. nigra, and M. rubra, based primarily on morphological characteristics like leaf shape and fruit type. This foundational treated Morus as a small group within the family, with seven species described overall, though only five are now recognized as valid mulberries. In the , Édouard Bureau expanded the in , revising Morus into five principal species (M. alba, M. celtidifolia, M. insignis, M. nigra, and M. rubra), along with 19 varieties and 11 subvarieties, emphasizing sectional divisions based on leaf morphology, such as entire versus lobed leaves, and pistillate features. Bureau's work marked a shift toward more detailed infrageneric groupings, influencing subsequent by incorporating variability in vegetative and reproductive structures. The 20th century saw significant revisions, with Gen'ichi Koidzumi's 1917 monograph proposing 24 species and one subspecies, organized into two major sections—Macromorus and Dolichostylae—primarily differentiated by style length in female flowers and syncarp shape. This proliferation reflected increased exploration of Asian diversity, but later systematists consolidated the count; for instance, Leroy in 1949 recognized 18 species across three subgenera (Eumorus, Gomphomorus, Afromorus), while Hotta in 1954 elevated it to 35 species. By the late 20th and early 21st centuries, syntheses like Chang et al. (1998) reduced the recognized species to 16, particularly for Chinese taxa, incorporating herbarium data and regional floras to address hybridization and synonymy issues. These consolidations, often to 10–16 species by the 2000s, highlighted the challenges of morphological plasticity in Morus, paving the way for molecular approaches. In the 2020s, phylogenomic studies have refined Morus using molecular markers, including nuclear ITS regions, trnL-trnF spacers, and full genomes, reclassifying the into approximately five sections such as Morus (encompassing Asian and North American lineages) and others like Dolichostylae, while resolving non-monophyly in earlier groupings. These analyses, such as & Ferguson (2012) and subsequent works, separated African and South American clades, with Hyb-Seq data from hundreds of nuclear loci revealing hybridization events and influencing biogeographic interpretations. A 2025 study analyzing complete genomes from 25 Morus accessions confirmed sectional boundaries through phylogenetic clustering into three major clades (wild, fruit, and leaf types), supporting M. alba and relatives as basal while highlighting intraspecific variation via simple sequence repeats. Additionally, a telomere-to-telomere assembly of wild M. mongolica in 2025 provided a high-quality reference, enabling precise delineation of chromosomal structures and further validation of phylogenetic relationships within the .

Species and Diversity

The genus Morus encompasses approximately 15–19 accepted , with taxonomic authorities varying in their counts; for example, a 2023 review estimates around 16 , while recognizes 17 as of 2024. Prominent include M. alba (white mulberry), M. nigra (black mulberry), M. rubra (red mulberry), M. laevigata, and M. mongolica, each distinguished by traits such as fruit color and morphology. Phylogenetic analyses based on nuclear ribosomal DNA (nrDNA) and chloroplast DNA (cpDNA) from 2025 studies reveal well-supported reflecting geographic origins and divergences. These include a primary Asian representing the genus's origin, contrasted with disjunct American (North and South) and an African lineage, with divergence times estimated around 38.67 million years ago for the . Hybridization contributes to phylogenetic complexity, notably the fertile hybrid M. × rubralba arising from M. alba and M. rubra, which exhibits asymmetrical favoring M. alba nuclear markers in up to 67% of hybrids. Diversity within Morus is evident in morphological traits like variable leaf lobing—from entire to deeply lobed—and fruit coloration ranging from white to black, alongside substantial . A 2025 nrDNA study of 542 accessions uncovered high intraspecific polymorphism, identifying 158 single-nucleotide polymorphisms (SNPs) and 15 insertions/deletions (InDels) that highlight evolutionary flexibility and support delimitation. Among recognized , M. alba is the most extensively cultivated worldwide, prized for its adaptability in and ornamental use. Conversely, M. mesozygia exemplifies conservation concerns in , classified as critically endangered in regions like due to habitat loss and limited distribution.

Biogeography and Ecology

Native and Introduced Ranges

The genus Morus encompasses 17 of trees and shrubs primarily native to temperate and subtropical regions of , with a concentration in and the Himalayan foothills where species such as M. alba, M. macroura, and others originated. Disjunct native distributions occur elsewhere, including M. rubra in eastern from southward to and westward to , M. nigra in southwestern (precise range obscured by ancient cultivation but centered in regions like ). Introduced ranges of Morus species, particularly M. alba, have expanded globally through human-mediated dispersal tied to , with the species now widespread in (introduced in the 1600s via colonial efforts to establish silk production), , (planted in since 1844 for mulberry cultivation), and (brought by Spanish colonizers post-conquest). In the United States, M. alba has become invasive across much of the lower 48 states except , aggressively colonizing disturbed areas, forest edges, and urban sites. It poses a significant threat in the eastern U.S. by outcompeting and hybridizing with the native M. rubra, potentially displacing it through faster maturation, prolific production, and transmission of root diseases. Phylogenetic analyses using genomes indicate a Central Asian cradle for the , with diversification driven by post-glacial migrations following the [Last Glacial Maximum](/page/Last Glacial Maximum), enabling northward and eastward expansions into temperate zones during warming. A 2025 biogeographic study of Morus cpGenomes from 25 accessions, including ancient trees in , confirms this Asian origin and highlights how climatic shifts facilitated the genus's radiation across before disjunct colonizations in other continents.

Habitat Preferences

Morus species thrive in temperate to subtropical climates, typically within USDA hardiness zones 4 to 9, where they exhibit tolerance to down to approximately -20°C and demonstrate resistance once established. These prefer full sun exposure but can adapt to partial shade, with optimal growth in regions receiving moderate annual rainfall of 400-1000 mm, though they endure both arid semi-desert conditions and occasional flooding in their native ranges. In terms of soil preferences, Morus favors well-drained loamy soils with a pH range of 5.5 to 7.0, showing adaptability to sandy, clayey, or even poor, infertile substrates as long as drainage is adequate. While tolerant of sporadic waterlogging, prolonged saturation leads to root damage and reduced vigor, underscoring their sensitivity to consistently wet conditions. Ecologically, Morus acts as a in disturbed habitats such as edges, old fields, and urban lots, rapidly colonizing gaps to stabilize and facilitate . As the primary host plant for silkworm larvae (), it supports ecosystems, while its fruits attract foraging birds and its canopy provides shade and nesting cover for . Morus forms symbiotic associations with soil microorganisms, including arbuscular mycorrhizal fungi that enhance nutrient uptake in nutrient-poor soils, and endophytic bacteria that promote growth and stress tolerance. In introduced regions, it often competes aggressively with native , sometimes acting as an invasive that displaces local species in open or riparian areas. In their native Asian ranges, Morus species show a particular affinity for riparian zones along rivers and streams in central and northern , where moist, alluvial soils support their establishment. Recent 2025 studies highlight Morus's high adaptability to , noting traits like and resilience that enable persistence across shifting agro-climatic zones.

Reproduction

Flowering and Pollination

Most Morus species are dioecious, though some can be monoecious, with flowers produced on separate individuals or occasionally the same . Male flowers form in staminate that produce , while female flowers develop in pistillate catkins bearing ovules. In the , flowering typically occurs in spring, from March to May, with blooms lasting about 1-2 weeks. Catkin sizes vary by species; for example, in red mulberry (), pistillate catkins measure 1-3 cm long, and staminate catkins reach 3-5 cm. Pollination in Morus is primarily anemophilous, relying on to transfer from to flowers. The dioecious nature necessitates proximity between and plants for successful set, as isolated females produce few or no viable fruits. exhibits high viability, often exceeding 89% in species like black mulberry (), supporting effective dispersal within local populations. Following , development proceeds rapidly, taking 4-8 weeks until ripening. Hybridization between phylogenetically distant Morus species, such as red mulberry (M. rubra) and white mulberry (M. alba), faces post-pollination barriers that reduce success rates and contribute to asymmetric gene flow. These barriers include pollen tube growth inhibition and embryo inviability, limiting interspecific crosses despite overlapping ranges.

Seed Dispersal and Propagation

Seed dispersal in Morus species primarily occurs through zoochory, where fruits are consumed by birds and mammals, facilitating the spread of via endozoochory. Birds such as and other frugivores play a key role in this process, ingesting the ripe, fleshy aggregate fruits and excreting viable seeds at distant locations, often promoting long-distance dispersal. Mammals, including red foxes, also contribute by eating the fruits and dispersing seeds through their scat, though birds are the dominant vector. In riparian habitats, some Morus species exhibit limited hydrochory, where seeds may float and travel short distances via water currents during floods, supplementing animal-mediated dispersal. This dispersal mechanism has enabled Morus to become invasive in introduced ranges, particularly M. alba, where bird-dispersed seeds establish dense thickets that outcompete native vegetation. Morus seeds are orthodox, tolerating and maintaining viability for 2-3 years under cool, dry storage conditions at 0-5°C. rates reach 70-90% following cold moist stratification for 30-100 days at 4-5°C, which breaks and enhances establishment, with optimal temperatures of 15-20°C post-stratification. Vegetative propagation is a prominent reproductive strategy in Morus, allowing clonal spread without . Root suckers emerge readily from injured or stems, enabling rapid colonization in disturbed soils and contributing to the persistence of wild populations. cuttings, taken in late spring, achieve rooting success rates of up to 80% when treated with auxins and maintained under , while cuttings root less reliably but are used for larger-scale . Layering, both ground and air methods, is effective for Morus, with branches rooting in moist soil or sphagnum moss, producing independent within one season. is rare in Morus, with most reproduction relying on sexual or vegetative means, though recent genomic analyses of wild populations indicate occasional clonal lineages maintained by vegetative sprouting.

Cultivation

History of Domestication

The domestication of Morus, particularly M. alba, began in central and northern around 2700 BCE, driven by its essential role in as the primary food source for silkworms (Bombyx mori) in production. This early cultivation marked a pivotal advancement in ancient , transforming mulberry from a wild species into a cornerstone of Chinese economy and . Archaeological from sites in and confirms practices dating to this period, with threads and cocoons indicating organized mulberry propagation. Ancient texts like the Shennong Bencao Jing, compiled around the 1st century CE but drawing on earlier traditions, further document mulberry's uses, classifying its leaves, fruits, and bark as therapeutic agents for ailments such as coughs and digestive issues while emphasizing its agricultural value. Mulberry cultivation disseminated westward along the trade network, with fabric reaching the by the 1st century BCE. However, and mulberry groves were introduced to the around 550 CE, integrating into Mediterranean societies for production. This expansion relied on M. alba's adaptability, allowing it to thrive in diverse climates from temperate to . By the early centuries CE, mulberry groves supported emerging in regions like Persia and , though the technology remained a closely guarded Chinese secret for millennia. European colonists introduced M. alba to the in the early 1600s, aiming to replicate Asian silk industries in the . In 1624, the mandated that every household plant at least four mulberry trees to bolster local , reflecting royal encouragement from King James I to reduce reliance on Asian imports. This effort extended to other colonies, including Georgia in 1733, where 500 trees were imported for experimental plantations. The witnessed intensified mulberry plantings in the United States, fueled by speculative booms in silk production, particularly with the introduction of fast-growing Morus multicaulis from around 1830. Prices for mulberry saplings surged dramatically, from $4 per hundred in 1834 to $30 per hundred by 1836, as states like promoted widespread cultivation through guidebooks and incentives. However, these trials collapsed by the 1840s due to mulberry blights, the labor-intensive nature of silkworm rearing, and inability to compete with inexpensive Chinese silk imports. Botanical studies during this era, including detailed species delineations, supported initial breeding efforts to enhance leaf yield and disease resistance for . In the , mulberry cultivation increasingly emphasized fruit varieties, with breeders in the United States and selecting for larger, sweeter drupes suitable for human consumption and ornamentals, diverging from sericulture dominance. This shift capitalized on M. alba and hybrids' native adaptability in Asian ranges, promoting sustainable amid declining silk interest.

Modern Practices

Modern mulberry cultivation emphasizes optimized and planting techniques to maximize productivity for both and production. Trees are typically spaced 5-10 meters apart to allow for canopy development and efficient harvesting, with provided regularly during phase to support root growth in the first 1-2 years. is conducted during in late winter to shape the tree, remove dead or diseased branches, and enhance yield by promoting new growth and improving air circulation. Varietal selection plays a crucial role in adapting mulberry to specific purposes and environmental challenges. For fruit production, hybrids such as 'Illinois Everbearing' (Morus alba × M. rubra) are favored for their extended ripening season and high-quality, sweet berries up to 4 cm long. Disease-resistant cultivars, including those bred to withstand bacterial caused by pv. mori, are increasingly selected to minimize losses in humid regions. Pest management in contemporary mulberry orchards relies on (IPM) strategies to control common threats like scale insects (e.g., white peach scale, Pseudaulacaspis pentagona) and mites, combining monitoring, cultural practices, and targeted interventions. In sustainable and organic farms, biological controls such as predatory insects and horticultural oils are prioritized over broad-spectrum chemicals to maintain ecosystem balance. Under optimal conditions, mature fruiting mulberry trees yield 10-20 kg of berries per tree annually, while sericulture-focused plantations achieve leaf production of 20-40 tons per per year through high-density planting and multiple harvests. dominates global , accounting for approximately 80% of worldwide production, which drives intensive mulberry cultivation advancements. Recent 2025 developments in genomic breeding have introduced drought-tolerant varieties, such as 'Guiyou 2024', identified through and analyses to enhance resilience in water-scarce areas without compromising yield.

Uses

Sericulture

, the practice of silk production, relies heavily on Morus species, particularly , as the primary host plant for the domestic silkworm . The leaves of M. alba serve as the exclusive feed for B. mori larvae, providing essential nutrients that enable the to spin cocoons composed of proteins. High-quality mulberry leaves typically contain 15-25% crude protein on a dry weight basis, which supports rapid larval growth and cocoon formation. Harvesting mulberry leaves for involves repeated to meet the silkworms' continuous demand for fresh foliage, with nutritional profiles optimized for larval development, including balanced and digestible carbohydrates. In tropical and subtropical regions, plants can yield 4-6 leaf crops per year through cycles spaced 10-12 weeks apart, allowing for sustainable production aligned with multiple silkworm rearing cycles. This intensive harvesting ensures leaves remain tender and nutrient-rich, as older leaves have reduced protein digestibility that can hinder silkworm health and silk yield. The global industry centers on Morus cultivation, with accounting for the majority of production, approximately 55% of the world's raw output as of 2023. Breeding programs focus on developing mulberry varieties with enhanced quality, such as higher protein content and disease resistance, to improve silkworm performance and overall silk efficiency. These efforts have sustained the industry's scale, where billions of B. mori cocoons are processed annually into silk threads. Silk from B. mori cocoons consists primarily of , the structural core protein comprising 70-80% of the fiber, coated by sericin, a hydrophilic gum-like protein making up 20-30%, which binds the filaments during spinning. Historically, maintained a near-monopoly on production for millennia, but this was effectively broken in the as expanded in , particularly in and , through technological advancements and established cultivation practices.

Culinary Applications

The fruits of Morus species, particularly M. alba and M. rubra, are widely consumed fresh for their sweet, juicy flavor, often harvested at peak ripeness from mid-June through August in temperate regions. These berries are versatile in preparations such as jams, jellies, pies, tarts, cakes, and breads, where their natural sweetness enhances baked goods and preserves. In Mediterranean cuisines, Morus nigra fruits are prized for desserts, including rich tarts and sorbets, due to their intense, dark berry taste. Processing methods extend the usability of mulberry fruits; they are frequently dried for long-term storage and used as a substitute in various recipes. produces mulberry wine, typically reaching 9-12% after primary and secondary stages, yielding a fruity, ruby-red beverage enjoyed in home and commercial settings. Regionally, in , mulberry pekmez—a thick molasses—is boiled down from the fruit juice and serves as a traditional in breakfast spreads, often mixed with or drizzled over pastries. Mulberry leaves contribute to culinary traditions, especially in , where young, tender leaves are blanched and incorporated into dishes like seasoned side salads or stir-fries with . In , they feature in rice preparations such as mulberry leaf —sushi-style rolls—or vegetable mixes, adding an earthy, nutty note when briefly cooked. Dried leaves are steeped to make a mild, caffeine-free , commonly consumed hot or iced in East Asian meals.

Medicinal and Pharmacological Uses

In traditional Chinese medicine, various parts of Morus species, particularly Morus alba and Morus nigra, have been utilized for their therapeutic properties. Mulberry leaves have been employed to alleviate symptoms of diabetes (known as "Xiao-ke"), hypertension, cough, sore throats, fever, and bronchitis, often through decoctions or teas that nourish the liver and improve vision while countering wind-related ailments. The root bark is valued for its cooling effects, helping to clear heat and treat conditions like diabetes mellitus by promoting diuresis and reducing inflammation. Bark and roots have also been traditionally used for their hypoglycemic effects, addressing weakness, fatigue, anemia, and kidney deficiencies by nourishing yin and blood. Pharmacological studies have substantiated many traditional uses, with a focus on antidiabetic and anti-inflammatory activities. Mulberry leaves contain and other antioxidants that exhibit anti-inflammatory effects by scavenging free radicals and modulating production. A key compound, 1-deoxynojirimycin (DNJ), acts as a potent α-glucosidase inhibitor, delaying digestion and reducing postprandial ; a 2025 review highlights DNJ's strong inhibitory potential against α-glucosidase and α-amylase, contributing to glycemic control. Additionally, a 2025 identifies over 50 bioactive compounds in , including , phenolic acids, and alkaloids, underscoring their pharmacological diversity. Clinical trials support the blood sugar-lowering effects of Morus extracts. In one randomized study, healthy adults consuming 280 mg of mulberry extract three times daily for 12 weeks experienced improved postprandial glycemic and insulinaemic responses to loads. Another trial involving individuals with impaired glucose tolerance showed that 1 g of mulberry taken three times daily after meals for 30 days reduced fasting blood glucose by 27%. Extracts rich in DNJ (12 mg three times daily before meals for 12 weeks) also modestly decreased serum triglycerides and improved profiles in subjects with borderline . For antimicrobial properties, Morus extracts demonstrate activity against bacteria such as Streptococcus mutans, Escherichia coli, Staphylococcus aureus, and periodontopathic species like Porphyromonas gingivalis, attributed to and phenolics that disrupt bacterial cell walls. Dosage guidelines for mulberry leaf extract in medicinal applications typically range from 1 to 3 g per day, divided into three doses taken before or after meals, with studies using 0.8–1 g three times daily for up to three months showing efficacy in glycemic control without significant adverse effects. Higher DNJ-enriched doses (e.g., 12–18 mg daily) have been tested for postprandial glucose attenuation, but consultation with healthcare providers is recommended to tailor usage.

Industrial Applications

Mulberry species, particularly , have been utilized for extracting natural pigments from their fruits, yielding deep dyes suitable for applications. The anthocyanin-rich extracts from black mulberry fruits provide vibrant coloration that has historically been applied to , , and fabrics, offering properties alongside aesthetic value. In traditional practices dating back to ancient , these fruit-based dyes were combined with mordants like to achieve fastness on natural fibers, contributing to durable and brownish hues in woven goods. Recent advancements emphasize their sustainability; as of 2025, encapsulated mulberry fruit extracts are being developed as eco-friendly alternatives to synthetic dyes in the , reducing environmental impact through biodegradable sourcing. The fibrous inner bark of Morus alba and related species has served as a raw material for high-quality paper production, particularly in East Asian traditions. In China, mulberry bark was processed into pulp for durable writing and printing papers as early as the Han dynasty, with the stems stripped, boiled, and beaten to yield fine, strong sheets. Modern adaptations include experimental papermaking from Morus indica twigs, where the bark is soaked and treated with sodium hydroxide to produce lightweight, white paper suitable for artisanal crafts. In the 19th century, United States agricultural experiments explored mulberry bark for paper amid broader silkworm cultivation efforts, though these trials focused more on fiber quality than large-scale production. Variants akin to Japanese washi, while primarily from paper mulberry (Broussonetia papyrifera), occasionally incorporate Morus bast fibers for enhanced texture in handmade sheets. Mulberry wood, derived from species like and , is a soft to medium valued for its workability in industrial woodworking. With an average dried of approximately 690 kg/m³, it offers a balance of strength and lightness, making it ideal for crafting furniture components, crates, and turned objects such as . The wood's fine grain and yellowish-to-brown heartwood allow easy and finishing, though its moderate (Janka rating around 1,680 lbf) suits indoor applications over heavy structural use. Beyond primary materials, mulberry waste supports production, enhancing industrial . Prunings and branches from plantations are pelletized into high-energy fuel, with calorific values comparable to conventional wood pellets, providing a source from agricultural byproducts. Extracts from mulberry roots and fruits are also incorporated into , where their tyrosinase-inhibiting compounds promote skin brightening and even tone in formulations like creams and serums. These applications underscore mulberry's versatility in eco-conscious industries as of 2025.

Nutrition and Bioactive Compounds

Nutritional Composition

The fruits of Morus species, commonly known as mulberries, are characterized by a high water content, typically comprising about 88% of their fresh weight, which contributes to their juicy texture and low caloric density of approximately 43 kcal per 100 g. Carbohydrates make up around 9.8 g per 100 g, primarily in the form of sugars such as glucose and , while protein content is modest at 1.4 g per 100 g and fat is negligible at 0.4 g per 100 g. is present at about 1.7 g per 100 g, aiding in digestive health. These values are based on aggregated data for raw mulberries from the (USDA) FoodData Central database. Micronutrients in mulberry fruits are notable, particularly at 36.4 mg per 100 g, representing a significant portion of the daily recommended intake, along with iron at 1.85 mg per 100 g. The skin of the contains , a , at concentrations up to 50.61 μg per g dry weight, varying by and ripeness. Other minerals include (194 mg per 100 g) and calcium (39 mg per 100 g), supporting balance and . These compositional elements highlight mulberries' role as a nutrient-dense, low-energy option. Mulberry leaves exhibit a higher protein content compared to the fruits, ranging from 15% to 31% on a dry weight basis across different genotypes, making them a valuable protein source in animal feeds and potential supplements. For instance, dried leaf powder from various cultivars contains 15.31% to 30.91% crude protein, alongside 27.6% to 36.66% . A 2025 study on 21 Turkish mulberry genotypes revealed significant nutritional variation, with showing the highest capacity at 47.68%, attributed to elevated levels of polyphenols and , while overall macronutrient profiles remained consistent with broader data. This intraspecific diversity underscores the adaptability of Morus for nutritional applications.
Nutrient (per 100 g raw fruit)AmountSource
88 gUSDA
Calories43 kcalUSDA
Carbohydrates9.8 gUSDA
Protein1.4 gUSDA
0.4 gUSDA
1.7 gUSDA
36.4 mgUSDA
Iron1.85 mgUSDA

Health Benefits and Pharmacology

Mulberry (Morus spp.) extracts, particularly from leaves, have demonstrated hypoglycemic effects primarily through the inhibition of α-glucosidase by 1-deoxynojirimycin (DNJ), a key iminosugar that delays carbohydrate digestion and reduces postprandial blood glucose spikes. Clinical trials indicate that doses of 1 g of mulberry leaf extract can lower glucose levels by approximately 12 mg/dL in healthy individuals, with more pronounced effects in type 2 diabetes patients, where postprandial glucose reductions of 1.04 mmol/L have been observed 30-90 minutes after ingestion. A 2022 systematic review of 14 randomized controlled trials confirmed that Morus alba supplementation significantly decreases postprandial glucose and insulin levels, supporting its role in diabetes management. Antioxidant properties of mulberry contribute to cardiovascular health by scavenging free radicals and mitigating , with leaf extracts exhibiting values of 8.5-16 μg/mL in and assays. These effects include reduced markers like (MDA) following 300 mg daily supplementation, alongside improvements in lipid profiles such as a 14% decrease in triglycerides. In animal models, mulberry polyphenols activate pathways like IRS-1/PI3K/AKT and AMPK, enhancing translocation for better and protecting vascular cells from oxidative damage. A 2022 of 21 studies further showed favorable impacts on cardiometabolic risk factors, including lowered LDL and . Anti-obesity benefits have been observed in models, where mulberry extracts at 300-500 mg/kg reduce body , visceral accumulation, and serum lipids by activating PPAR-γ receptors, which regulate and . This mechanism involves downregulation of proinflammatory cytokines like TNF-α and IL-6, promoting . A 2025 comprehensive review synthesizing over 190 studies highlighted consistent efficacy across antidiabetic, , and anti-obesity outcomes, with mulberry extracts showing up to 20% reductions in blood glucose in diabetic models. Pharmacologically, mulberry's safety profile in supplements is favorable, with no or reported in human and animal studies at typical doses of 280-1000 mg/day; however, potential interactions include additive hypoglycemic effects with antidiabetes drugs and modulation of /P-gp, which may alter bioavailability of substrates like cyclosporine. A meta-analysis of 13 trials reinforced these benefits for blood glucose control (weighted mean difference of -18.06 mg/dL) while noting the need for monitoring in .

Safety Concerns

Toxicity

The unripe fruits of Morus species contain a milky sap that acts as a mild irritant, potentially causing , stomach upset, and hallucinations if ingested. This is present throughout the except in ripe , and contact with it can also irritate the skin. The root bark of contains , which in excess can contribute to effects and digestive disturbances due to their anti-nutritional properties. Overconsumption of mulberry fruits or leaves may lead to digestive upset, including , owing to their properties. Additionally, rare traces of (1.01–2.14 mg/kg) have been detected in mulberry leaves, though seeds show no confirmed presence, posing minimal at typical intake levels. Mulberry leaves are and commonly fed to silkworms without issues. Toxicity studies on Morus alba extracts in rodents indicate low risk, with an LD50 exceeding 5 g/kg body weight and no observed adverse effects at doses up to 7.5 g/kg/day. No human fatalities from Morus consumption have been reported, reflecting its overall mild toxicity profile.

Allergenicity

Morus alba pollen is a significant source of respiratory allergies, including allergic rhinitis and asthma, particularly during the spring pollination season when it triggers hay fever symptoms such as sneezing, nasal congestion, and itchy eyes. This high allergenicity is attributed to key proteins in the pollen, including profilins (14-15 kDa), which act as pan-allergens and contribute to cross-reactivity with birch pollen (Bet v 2) and other tree pollens like olive and wall pellitory. Sensitization to Morus pollen shows variable prevalence, with rates of 1.4% to 7% reported among allergic patients in urban areas of the and where the trees are commonly planted for landscaping or . In regions like and northern , sensitization studies indicate higher exposure risks in urban settings, affecting 5-10% of pollen-allergic individuals during peak seasons ( to ), often co-occurring with sensitivities to multiple aeroallergens. Beyond inhalation, direct contact with Morus sap can induce in sensitive individuals, manifesting as irritation, redness, or urticaria due to irritant components. Airborne contact reactions from or sap vapors have also been documented, particularly in occupational settings like or harvesting. Management of Morus-related primarily involves avoidance strategies, such as limiting outdoor activities during peak release in spring and removing trees from high-exposure urban sites, alongside symptomatic relief with antihistamines or nasal corticosteroids; desensitization through specific is effective for pollen-sensitized patients in severe cases.

Cultural Significance

Symbolism and Folklore

In Chinese lore, the mulberry tree symbolizes and , often linked to its essential role in , where its leaves nourish silkworms that produce , a material revered as a divine gift associated with eternal life. The tree's representation as the "" or "herb of immortality" stems from ancient beliefs in its life-sustaining properties, including its use in herbal remedies and its mythological ties to longevity in Taoist traditions. Among Native American communities, the red mulberry () served as a reliable source of nutritious fruit, dye, and fiber, with groups like the using it for food such as dumplings and jams, as well as in medicinal preparations for ailments like . Greek folklore features the mulberry in the myth of , where the tragic lovers' blood stains the tree's white berries red, transforming the fruit into an enduring emblem of forbidden and sorrow rather than the lovers themselves becoming trees. In ancient , the mulberry was associated with worship, deified as a of and . In , festivals at shrines like Kokage-San Jinja involve prayers for silkworm health, with offerings of mulberry leaves and cocoons held biannually—on March 28 for successful cultivation and October 23 for thanksgiving—to honor the tree's vital connection to and communal well-being. Biblical references to mulberries appear in strategic and metaphorical contexts, such as in 2 Samuel 5:23-24 and 1 Chronicles 14:14-15, where the sound of movement in the mulberry (or ) trees signals divine timing for David's battle against the . In the , Luke 17:6 uses the mulberry tree to illustrate the power of even small faith, capable of uprooting it and planting it in the sea. In 19th-century European superstitions, the mulberry's deep red fruit staining was often viewed as an ill omen, evoking associations with blood and tragedy from persisting Greek myths, while German folklore linked the berries to evil, claiming the devil polished his boots with the tree's roots.

Representation in Art and Literature

In classical Western literature, the mulberry tree (Morus) features prominently in Ovid's Metamorphoses (ca. 8 CE), particularly in the tragic tale of Pyramus and Thisbe, where the lovers arrange to meet beneath a mulberry tree whose white berries are stained red by Pyramus's blood after a fatal misunderstanding, permanently altering the fruit's color to symbolize eternal mourning and the transformative power of love and death. This motif of bloodied mulberries recurs in Willa Cather's O Pioneers! (1913), where a white mulberry tree in Marie Shabata's orchard serves as the site of her murder alongside lover Emil Bergson, with the red-stained berries evoking Ovid's myth while underscoring themes of passion's peril and the harsh realities of American frontier life. In ancient , mulberry trees are among the most frequently invoked plants in the Shijing (Book of Songs, ca. 11th–7th centuries BCE), the oldest extant collection of , where they symbolize grace, abundance, and seasonal renewal; for instance, poems describe the tree's tender leaves in lowlands as metaphors for delight in reunion or the rhythms of agrarian existence tied to . The tree's cultural resonance extends to later works, such as those exploring production's societal role, reflecting Morus alba's foundational place in East Asian poetic traditions. In visual art, captured the mulberry tree's vibrant resilience in his 1889 oil painting The Mulberry Tree, executed at the Saint-Rémy asylum with bold, swirling brushstrokes depicting golden autumn foliage against a turbulent blue sky, which he deemed one of his most assured landscapes amid personal turmoil. In Chinese artistic traditions, mulberries appear in functional yet aesthetically refined depictions of daily labor, as in Wu Jun's late 19th-century ink and watercolor Gathering 'Jingsang' Mulberry Leaves from a album on the silk industry, showing workers scaling ladders to harvest leaves from tall Morus trees, emphasizing the plant's economic centrality in imperial society. The Shakespearean mulberry—a legendary tree purportedly planted by the playwright in his garden around 1600—has inspired subsequent artworks and relics, including 18th-century wood carvings from its felled trunk, symbolizing literary heritage through its grafted connection to the bard's persona.

References

  1. https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:30004492-2
  2. https://www.treesandshrubsonline.org/articles/morus/
  3. https://www.mdpi.com/2073-4395/13/8/2021
  4. https://www.ijplantenviro.com/index.php/IJPE/article/download/1221/666
  5. https://pmc.ncbi.nlm.nih.gov/articles/PMC8241616/
  6. https://www.sciencedirect.com/science/article/pii/S266671932030011X
  7. https://www.invasive.org/species/subject.cfm?sub=6050
  8. https://www.feedipedia.org/node/123
  9. https://pmc.ncbi.nlm.nih.gov/articles/PMC10650418/
  10. https://gobotany.nativeplanttrust.org/species/morus/alba/
  11. https://www.mdpi.com/2311-7524/8/12/1140
  12. https://pmc.ncbi.nlm.nih.gov/articles/PMC8100788/
  13. https://www.fs.usda.gov/database/feis/plants/tree/moralb/all.html
  14. https://www.fs.usda.gov/rm/pubs_series/wo/wo_ah727/wo_ah727_728_732.pdf
  15. https://plants.ces.ncsu.edu/plants/morus-rubra/
  16. http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=121220
  17. https://www.sciencedirect.com/science/article/abs/pii/S105579032400085X
  18. https://www.researchgate.net/publication/351238274_Systematics_of_the_Genus_Morus_L_Moraceae
  19. https://bioone.org/journals/systematic-botany/volume-37/issue-2/036364412X635485/Phylogenetics-of-Morus-Moraceae-Inferred-from-ITS-and-trnL-trnF/10.1600/036364412X635485.full
  20. https://bmcgenomics.biomedcentral.com/articles/10.1186/s12864-025-11823-3
  21. https://www.nature.com/articles/s41597-025-05040-6
  22. https://www.mdpi.com/2223-7747/14/16/2570
  23. https://www.nature.com/articles/s41598-025-97618-8
  24. https://pubmed.ncbi.nlm.nih.gov/16156816/
  25. https://www.zimbabweflora.co.zw/speciesdata/species.php?species_id=120130
  26. https://www.fs.usda.gov/nsl/Wpsm/Morus.pdf
  27. https://tropical.theferns.info/viewtropical.php?id=Morus%2Bmacroura
  28. https://www.wildflower.org/plants/result.php?id_plant=moru2
  29. https://www.ontarioinvasiveplants.ca/wp-content/uploads/2025/07/OIPC_BMP_WhiteMulberry_April2021_D6_WEB.pdf
  30. https://pir.sa.gov.au/aghistory/industries/other_industries/silk
  31. https://www.invasiveplantatlas.org/subject.cfm?sub=6050
  32. https://www.invasive.org/alien/pubs/midatlantic/moal.htm
  33. https://www.wildabundance.net/blog/mulberry-trees/
  34. https://toptropicals.com/catalog/image/6/morus_hybrid.htm
  35. https://uncommonfruit.cias.wisc.edu/mulberry/
  36. https://raintreenursery.com/pages/growing-fruit-trees-mulberries
  37. https://www.nature.com/articles/s41598-024-76455-1
  38. https://thepolycultureproject.medium.com/mo-mulberry-the-essential-guide-to-all-you-need-to-know-about-mulberry-28a0c11b611
  39. https://pubmed.ncbi.nlm.nih.gov/30047827/
  40. https://pmc.ncbi.nlm.nih.gov/articles/PMC4923160/
  41. https://www.frontiersin.org/journals/microbiology/articles/10.3389/fmicb.2025.1618773/full
  42. https://www.inaturalist.org/guide_taxa/882846
  43. https://www.sciencedirect.com/science/article/pii/S2665972723000545
  44. https://www.sciencedirect.com/science/article/abs/pii/S0377840125003530
  45. https://link.springer.com/article/10.1007/s12600-025-01261-y
  46. https://weedguide.cfaes.osu.edu/singlerecord.asp?id=77
  47. https://growplant.org/blog/mulberry-tree-life-cycle/
  48. https://research.fs.usda.gov/treesearch/download/32653.pdf
  49. https://pmc.ncbi.nlm.nih.gov/articles/PMC11351111/
  50. https://www.researchgate.net/publication/393172421_Pollen_Performance_and_Morphology_of_Black_Mulberry_Morus_nigra_L_Genotypes
  51. https://bsapubs.onlinelibrary.wiley.com/doi/10.3732/ajb.1200085
  52. https://www.jstor.org/stable/1938870
  53. https://www.cabidigitallibrary.org/doi/full/10.1079/cabicompendium.34816
  54. http://scielo.sld.cu/scielo.php?script=sci_arttext&pid=S0864-03942017000400003&lng=en&tlng=en
  55. https://abcbot.pl/pdf/47_2/023-26.pdf
  56. https://www.researchgate.net/publication/375497073_Mulberry_cutting_propagation_A_Review
  57. https://mulberrytrees.co.uk/vegetative/
  58. https://www.researchgate.net/publication/390741215_GENETICS_CYTOGENETICS_AND_GENOMICS_OF_MULBERRY_MORUS_SPP
  59. https://www.treesandshrubsonline.org/articles/morus/morus-alba/
  60. https://phl.juncture-digital.io/mulberry
  61. https://www.worldscientific.com/doi/10.1142/S0192415X23500775
  62. https://en.unesco.org/silkroad/about-silk-roads
  63. https://www.fao.org/4/x9895E/x9895e02.htm
  64. https://www.china-silkroad-travel.com/silk-road-culture/685.html
  65. https://hgic.clemson.edu/red-vs-white-mulberry-morus-rubra-vs-morus-alba-in-south-carolina/
  66. https://history.textiles.ncsu.edu/textile-industry-history/silk-in-america/
  67. https://connecticuthistory.org/connecticuts-mulberry-craze/
  68. https://digitalcommons.lindenwood.edu/cgi/viewcontent.cgi?article=1043&context=student-research-papers
  69. https://www.pubhort.org/members/showdocument?tarball=1&series=aps&vid=41&iid=1&article=2&layout=pubhort&action=showdocument&lidvan=aps
  70. https://www.starkbros.com/growing-guide/how-to-grow/fruit-trees/mulberry-trees/planting
  71. https://www.starkbros.com/growing-guide/how-to-grow/fruit-trees/mulberry-trees/pruning
  72. https://onegreenworld.com/product/illinois-everbearing-mulberry/
  73. https://pnwhandbooks.org/plantdisease/host-disease/mulberry-morus-spp-bacterial-blight
  74. https://ipm.ucanr.edu/PMG/PESTNOTES/pn7408.html
  75. https://paradisenursery.com/pages/pests-mulberry-trees
  76. https://www.apnikheti.com/en/pn/agriculture/horticulture/fruit/mulberry
  77. https://www.fao.org/4/x9895e/x9895e04.htm
  78. https://sino-silk.com/top-silk-producing-countries/
  79. https://ietresearch.onlinelibrary.wiley.com/doi/abs/10.1049/syb2.70004
  80. https://agriculture.institute/host-plant-cultivation/mulberry-leaf-yield-quality-india/
  81. https://www.fao.org/4/x9895e/x9895e0t.htm
  82. https://journalacri.com/index.php/ACRI/article/download/1418/3443/1989
  83. https://textileexchange.org/app/uploads/2024/09/Materials-Market-Report-2024.pdf
  84. https://www.mdpi.com/2079-4983/15/11/322
  85. https://www.silkroad.travel/interesting-facts-about-silk/
  86. https://www.starkbros.com/growing-guide/how-to-grow/fruit-trees/mulberry-trees/harvesting
  87. https://www.hava-baemek.co.il/en/tour/mulberry
  88. https://www.sciencedirect.com/science/article/pii/S0023643823001871
  89. https://gastronomica.org/2013/10/31/slowing-down-for-turkish-mulberry-molasses/
  90. https://pmc.ncbi.nlm.nih.gov/articles/PMC10650630/
  91. https://www.wonju.go.kr/english/contents.do?key=2953&
  92. https://www.rishi-tea.com/blogs/journal/mulberry-leaf
  93. https://www.nccih.nih.gov/health/white-mulberry-leaf
  94. https://www.vinmec.com/eng/blog/mulberry-leaves-uses-and-benefits-en
  95. https://www.sciencedirect.com/science/article/pii/S1756464623002190
  96. http://www.itmonline.org/arts/morus.htm
  97. https://www.researchgate.net/publication/369867820_Medicinal_values_of_mulberry_Morus_spp_A_review
  98. https://pubs.rsc.org/en/content/articlehtml/2025/fo/d4fo04954c
  99. https://www.researchgate.net/publication/388128757_A_Systematic_review_on_the_phytochemistry_isolated_compounds_nutritional_benefits_pharmacology_and_toxicology_of_the_plant_species_Morus_alba_L
  100. https://pmc.ncbi.nlm.nih.gov/articles/PMC8047566/
  101. https://www.drugs.com/npp/white-mulberry.html
  102. https://pmc.ncbi.nlm.nih.gov/articles/PMC2935155/
  103. https://www.nature.com/articles/s41598-025-18872-4
  104. https://pmc.ncbi.nlm.nih.gov/articles/PMC6412198/
  105. https://www.webmd.com/vitamins/ai/ingredientmono-1250/white-mulberry
  106. https://www.sciencedirect.com/science/article/abs/pii/S0965229919317315
  107. https://www.researchgate.net/publication/274638827_Black_mulberries_Morus_nigra_as_a_natural_dye_for_animal_tissues_staining
  108. https://bioticapublications.com/journal-backend/articlePdf/696c265288.pdf
  109. https://www.scirp.org/journal/paperinformation?paperid=77851
  110. https://www.researchgate.net/publication/372262926_Functional_Food_Development_of_Natural_Colorant_Additives_from_Mulberry_Fruit_Extract_Morus_nigra_L_with_Bioactive_and_Hedonic_Evaluation_Natural_Food_Colorant_from_Mulberry_Fruit
  111. https://smallbatchjamco.com/blogs/news/the-world-of-the-mulberry-silk-stationery-to-sweets
  112. https://www.researchgate.net/publication/388498817_EXTRACTION_OF_PAPER_FROM_MULBERRY_MORUS_INDICA_L_TWIGS_AND_ITS_ANALYSIS
  113. https://groups.io/g/Papermaking/topic/white_mulberry_for_paper/56576554
  114. https://www.wood-database.com/mulberry/
  115. https://www.woodworkingnetwork.com/wood/wood-explorer/red-mulberry-morus-rubra
  116. https://www.mdpi.com/1996-1073/18/13/3541
  117. https://www.makingcosmetics.com/BOT-MULBE-01.html?lang=en_US
  118. https://skintypesolutions.com/blogs/skincare/mulberry-extract
  119. https://www.agronomyjournals.com/archives/2025/vol8issue2/PartC/8-2-27-469.pdf
  120. https://www.growables.org/information/TropicalFruit/documents/MulberryNutrient.pdf
  121. https://pubmed.ncbi.nlm.nih.gov/25593373/
  122. https://pubmed.ncbi.nlm.nih.gov/17135021/
  123. https://pubmed.ncbi.nlm.nih.gov/40603370/
  124. https://pmc.ncbi.nlm.nih.gov/articles/PMC12381166/
  125. https://doi.org/10.1016/j.fbio.2024.104218
  126. https://doi.org/10.3390/nu17142262
  127. https://pmc.ncbi.nlm.nih.gov/articles/PMC9152381/
  128. https://japsonline.com/admin/php/uploads/3773_pdf.pdf
  129. https://pubmed.ncbi.nlm.nih.gov/26972284/
  130. https://www.researchgate.net/publication/22150678_Pharmacological_studies_on_root_bark_of_mulberry_tree_Morus_alba_L
  131. https://pmc.ncbi.nlm.nih.gov/articles/PMC11121110/
  132. https://m.facebook.com/SymbiosisInt/photos/did-you-know-that-mulberry-leaves-relieve-bloating-in-cows-no-neither-did-i-but-/848724974146468/
  133. https://www.sciencedirect.com/science/article/abs/pii/S027323001830076X
  134. https://pmc.ncbi.nlm.nih.gov/articles/PMC7029495/
  135. https://www.thermofisher.com/phadia/us/en/resources/allergen-encyclopedia/t70.html
  136. https://plants.ces.ncsu.edu/plants/morus-alba/
  137. https://www.thermofisher.com/allergy/us/en/allergen-fact-sheets/mulberry-tree.html
  138. https://greensilk.com/history-mulberry-leaf-history/
  139. https://vineyardgazette.com/news/2020/06/30/round-mulberry-bush
  140. https://albuquerqueherbalism.com/2025/04/20/the-versatile-mulberry/
  141. https://mcclungmuseum.utk.edu/2022/06/27/plantofthemonth-mulberry-morus-rubra/
  142. https://pressbooks.pub/iagtm/chapter/582/
  143. https://www.researchgate.net/publication/378119857_Studies_on_the_Origin_and_Cultural_Value_of_Chinese_Mulberry_Culture
  144. https://tsukublog.wordpress.com/2024/10/20/ancient-rituals-for-silkworms-at-kokage-san-jinja-hanging-on-to-life-by-a-very-fine-silk-thread-october-23rd-and-march-28th/
  145. https://biblehub.com/topical/m/mulberry-tree.htm
  146. https://www.intothewaters.com/blog/what-did-jesus-mean-that-a-mustard-seed-uproots-the-mulberry-tree
  147. https://bgdailynews.com/2022/07/24/the-folklore-of-plants-mulberry/
  148. https://www.cambridge.org/core/journals/ramus/article/red-and-white-in-ovids-metamorphoses-the-mulberry-tree-in-the-tale-of-pyramus-and-thisbe/4DF708415FF33819D8C25F48BB1FE3AC
  149. https://cather.unl.edu/scholarship/catherstudies/12/cs012.tsank
  150. https://oxfordre.com/literature/display/10.1093/acrefore/9780190201098.001.0001/acrefore-9780190201098-e-1267
  151. https://www.nortonsimon.org/art/detail/M.1976.09.P/
  152. https://collections.vam.ac.uk/item/O66947/gathering-jingsang-mulberry-leaves-painting-wu-jun/
  153. https://journal.transformativeworks.org/index.php/twc/article/view/2619
Add your contribution
Related Hubs
User Avatar
No comments yet.