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Ginkgo biloba, commonly known as ginkgo ( ), also known as the maidenhair tree, and often misspelled "gingko" (but see #Etymology below) is a species of gymnosperm tree native to East Asia. It is the last living species in the order Ginkgoales, which first appeared over 290 million years ago. Fossil similar to the living species, belonging to the genus Ginkgo, extend back to the Middle Jurassic epoch approximately 170 million years ago. The tree was cultivated early in human history, remains commonly planted, and is widely regarded as a living fossil.
G. biloba is a long-lived, disease-resistant, Dioecy tree with unique fan-shaped Leaf, capable of Cloning, and known for its striking yellow autumn foliage and resilience in disturbed environments. It was known historically as "silver fruit" or "white fruit" in Chinese language and called “ginkgo” due to a centuries-old transcription error. It is closely related to Cycad and characterized by unique Seed that resemble Apricot but are not true Fruit.
G. biloba, once widespread but thought Extinction in the wild for centuries, is now commonly cultivated in East Asia, with some genetically diverse populations possibly representing rare wild survivors in southwestern China’s mountainous regions. Some G. biloba trees have survived extreme events like the Hiroshima atomic bomb and others showcasing extreme longevity; G. biloba specimens have been measured in excess of 1,600 years, and the largest living trees are estimated to exceed 3,500 years. Today it is widely planted in City worldwide for its pollution tolerance and ornamental value.
G. biloba can pose health risks including potential Carcinogen, Allergy, poisoning from seeds due to ginkgotoxin, Drug interaction, and Adverse effect such as bleeding and Neurology symptoms, especially with excessive or improper use. G. biloba wood is valued for its durability and used in Craft and sake-making, while its seeds are popular in Asian cuisine despite health risks. While widely marketed for Nootropic benefits, clinical research shows limited medical effectiveness except possibly for dementia, with approval in the European Union but not by the United States Food and Drug Administration.
The species is heterophyllous (two types of leaves); those on the long shoots are thicker, have higher rates of photosynthesis, higher vein density and leaf hydraulic conductance, while those on the short shoots are better at handling drought. Hydraulic and Economical Traits in Short- and Long-Shoot Leaves of Ginkgo biloba Males and Females Leaves of long shoots are also usually notched or lobed, but only from the outer surface, between the veins. They are borne both on the more rapidly growing branch tips, where they are alternate and spaced out, and also on the short, stubby spur shoots, where they are clustered at the tips. Leaves are green both on the top and bottom and have stomata on both sides. During autumn, the leaves turn a bright yellow and then fall, sometimes within a short space of time (one to fifteen days). Leaves of the cultivar 'Tubifolia' have funnel-shaped leaves.
Ginkgo prefers full sun and grows best in environments that are well-watered and well-drained. The species shows a preference for disturbed sites; in the "semiwild" stands at , many specimens are found along stream banks, rocky slopes, and cliff edges. Accordingly, ginkgo retains a prodigious capacity for vegetative growth. It is capable of sprouting from embedded buds near the base of the trunk (, or basal chichi) in response to disturbances, such as soil erosion. Old specimens are also capable of producing aerial roots on the undersides of large branches in response to disturbances such as crown damage; these roots can lead to successful clonal reproduction upon contacting the soil. These strategies are evidently important in the persistence of ginkgo; in a survey of the "semiwild" stands remaining in Tianmu Mountain, 40% of the specimens surveyed were multi-stemmed, and few saplings were present.
Female plants do not produce cones. Two are formed at the end of a stalk, and after anemophily, one or both develop into fruit-like structures containing seeds. The fruits are 1.5–2 cm long, with a soft, fleshy, yellow-brown outer layer (the sarcotesta) that is attractive in appearance, but contains butyric acid (also known as butanoic acid) and smells foul like rancidification butter or vomit when fallen. Beneath the sarcotesta is the hard sclerotesta (the "shell" of the seed) and a papery endotesta, with the nucellus surrounding the female gametophyte at the center.
The fertilization of ginkgo seeds occurs via motility sperm, as in cycads, ferns, mosses, and algae. The sperm are large (about 70–90 micrometres) and are similar to the sperm of cycads, which are slightly larger. Ginkgo sperm were first discovered by the Japanese botanist Sakugoro Hirase in 1896. The sperm have a complex multi-layered structure, which is a continuous belt of basal bodies that form the base of several thousand flagella which have a cilia-like motion. The flagella/cilia apparatus pulls the body of the sperm forwards. The sperm have only a tiny distance to travel to the archegonia, of which there are usually two or three. Two sperm are produced, one of which successfully fertilizes the ovule. Fertilization of ginkgo seeds occurs just before or after they fall in early autumn. Embryos may develop in the seeds before or after they drop from the tree.
In 2020, a study in China of ginkgo trees up to 667 years old showed little effects of aging, finding that the trees continued to grow with age and displayed no genetic evidence of senescence, and continued to make indefinitely.
The scientific name Ginkgo is the result of a spelling error that occurred three centuries ago. Kanji typically have multiple pronunciations in Japanese, and the characters 銀杏 used for ginnan can also be pronounced ginkyō. Engelbert Kaempfer, the first Western world to investigate the species in 1690, wrote down this pronunciation in the notes that he later used for the Amoenitates Exoticae (1712) with the "awkward" spelling "ginkgo". This appears to be a simple error of Kaempfer; taking his spelling of other Japanese words containing the syllable "kyō" into account, a more precise romanization following his writing habits would have been "ginkio" or "ginkjo". Linnaeus, who relied on Kaempfer when dealing with Japanese plants, adopted the spelling given in Kaempfer's "Flora Japonica" ( Amoenitates Exoticae, p. 811). Kaempfer's drawing can be found in Hori's article.
The ginkgo is classified in its own division, the Ginkgophyta, comprising the single class Ginkgoopsida, order Ginkgoales, family Ginkgoaceae, genus Ginkgo and is the only extant taxon within this group. It is one of the best-known examples of a living fossil, because Ginkgoales other than G. biloba are not known from the fossil record after the Pliocene.
Such plants with leaves that have more than four veins per segment have customarily been assigned to the taxon Ginkgo, while the taxon Baiera is used to classify those with fewer than four veins per segment. Sphenobaiera has been used for plants with a broadly wedge-shaped leaf that lacks a distinct leaf stem.
The Ginkgophyta declined in diversity as the Cretaceous progressed, and by the Paleocene, Ginkgo adiantoides was the only Ginkgo species left in the Northern Hemisphere, while a markedly different (and poorly documented) form persisted in the Southern Hemisphere. Along with that of ferns, cycads, and cycadeoids, the species diversity in the genus Ginkgo drops through the Cretaceous, at the same time the flowering plants were on the rise; this supports the hypothesis that, over time, flowering plants with better adaptations to disturbance displaced Ginkgo and its associates.
At the end of the Pliocene, Ginkgo fossils disappeared from the fossil record everywhere except in a small area of central China, where the modern species survived.
At least morphologically, G. gardneri and the Southern Hemisphere species are the only known post-Jurassic taxa that can be unequivocally recognised. The remainder may have been or subspecies. The implications would be that G. biloba had occurred over an extremely wide range, had remarkable genetic flexibility and, though evolution genetically, never showed much speciation.
While it may seem improbable that a single species may exist as a contiguous entity for many millions of years, many of the ginkgo's life-history parameters fit: Extreme longevity; slow reproduction rate; (in Cenozoic and later times) a wide, apparently contiguous, but steadily contracting distribution; and (as far as can be demonstrated from the fossil record) extreme ecological conservatism (restriction to disturbed streamside environments).
Modern-day G. biloba grows best in environments that are well-watered and drained, and the extremely similar fossil Ginkgo favored similar environments: The sediment record at the majority of fossil Ginkgo localities indicates it grew primarily in disturbed environments, such as along streams. Ginkgo, therefore, presents an "ecological paradox" because while it possesses some favorable traits for living in disturbed environments (clonal reproduction) many of its other life-history traits are the opposite of those exhibited by modern plants that thrive in disturbed settings (slow growth, large seed size, late reproductive maturity).
The genus name is regarded as a misspelling of the Japanese pronunciation () for the kanji meaning "silver apricot", which is found in Chinese herbology literature such as 日用本草 (Daily Use Materia Medica) (1329) and Compendium of Materia Medica 本草綱目 published in 1578.T. Hori, A historical survey of Ginkgo biloba based on Japanese and Chinese classical literatures, Plant Morphology, 2001, 31, 31–40
Despite its spelling, which is due to a complicated etymology including a transcription error, "ginkgo" is usually pronounced , which has given rise to the common alternative spelling "gingko". The spelling pronunciation is also documented in some dictionaries.
Engelbert Kaempfer first introduced the spelling ginkgo in his book of 1712. (with illustration) It is considered that he may have misspelled "Ginkjo" or "Ginkio" (both consistent with his treatment of Japanese in the same work) as "Ginkgo". This misspelling was included by Linnaeus in his book See page 131 of Car. a Linné Mantissa plantarum: Generum editionis VI. et specierum editionis II, available at [3] and has become the name of the tree's genus. The specific epithet is New Latin for "two-lobed".
For centuries, it was thought to be extinct in the wild, but is now a common tree cultivated throughout eastern China, Korea, and Japan. Many municipalities in Korea and Japan use Ginkgos as street trees, and Ginkgo leaves are the emblem of prominent educational institutions such as the University of Tokyo and Sungkyunkwan University in South Korea. Despite their widespread habitat, high genetics exists among ginkgo trees, with some Chinese scholars suggesting that ginkgo trees in these areas may have been planted and preserved by Chinese monks over about 1,000 years. A study demonstrates a greater genetic diversity in Southwestern China populations, supporting glacial refugia in mountains surrounding the eastern Tibetan Plateau, where several old-growth candidates for wild populations have been reported. Whether native ginkgo populations still exist has not been demonstrated unequivocally, but there is genetic evidence that these Southwestern populations may be wild, as well as evidence that the largest and oldest G. biloba trees may be older than surrounding human settlements.
Where it occurs in the wild, Ginkgo is found infrequently in deciduous forests and valleys on acidic loess (i.e. fine, silty soil) with good drainage. The soil it inhabits is typically in the pH range of 5.0 to 5.5.
G. biloba is also commonly manually planted in cities across the United States and Europe. This species is highly tolerant to pollution and serves as a visually appealing, shade-providing tree in many cities and gardens.
Many intentionally planted ginkgos are male grafted onto plants propagated from seed, because the male trees will not produce the malodorous seeds. The popular cultivar 'Autumn Gold' is a clone of a male plant.
The disadvantage of male Ginkgo biloba trees is that their pollen is highly allergenic. They have an OPALS (Ogren Plant Allergy Scale) rating of 7 (out of 10), whereas female trees, which can produce no pollen, have an OPALS allergy scale rating of 2.
Female cultivars include 'Liberty Splendor', 'Santa Cruz', and 'Golden Girl', the latter so named because of the striking yellow color of its leaves in the fall; all female cultivars release zero pollen.
Many cultivars are listed in the literature in the UK, of which the compact 'Troll' has gained the Royal Horticultural Society's Award of Garden Merit.
Ginkgos adapt well to the urban environment, tolerating pollution and confined soil spaces. They rarely have disease problems, even in urban conditions, and are attacked by few insects.
Ginkgos are popular subjects for growing as miniature landscapes known as penjing and bonsai; they can be kept artificially small and tended over centuries. The trees are easy to propagate from seed.
The six trees are still alive: They are marked with signs at 報専坊 temple (planted in 1850), Shukkei-en (planted about 1740), Jōsei-ji (planted 1900), at the former site of Senda Elementary School near Miyukibashi, at the Myōjōin temple, and an Edo period-cutting at Anraku-ji temple.
Modern scholarship has established that ginkgos arrived from China in the 14th century, and a 1990 tree-ring measurement indicated the kakure-ichō's age to be about 500 years.
On 10 March 2010, the tree blew down in a storm, but the stump has since sprouted vigorously.
When eaten in large quantities or over a long period, the seeds may cause poisoning by ginkgotoxin (4'-O-methylpyridoxine, MPN), as found in a few . A heat-stable compound not destroyed by cooking, MPN may cause convulsions, which were alleviated by treatment with pyridoxine phosphate (vitamin B6), according to limited studies.
Some people are sensitive to the chemicals in the sarcotesta, the outer fleshy coating. These people should handle the seeds with care when preparing the seeds for consumption, wearing disposable gloves. The symptoms are allergic contact dermatitis, or similar to that caused by contact with poison ivy.
of using ginkgo supplements may include increased risk of bleeding, gastrointestinal discomfort, nausea, vomiting, diarrhea, headaches, dizziness, heart , and restlessness. Although use of standardized Ginkgo biloba leaf extracts in moderate amounts appears to be safe, excessive use may have undesirable effects, especially in terms of . The dosing of , such as warfarin or antiplatelet medication, may be adversely affected by using ginkgo supplements.
According to a systemic review, the effects of ginkgo on pregnant women may include increased bleeding time, and there is inadequate information about safety during lactation.
Ginkgo pollen may produce allergic reactions. Ginkgo biloba leaves and sarcotesta contain ginkgolic acids which are highly long-chain alkylphenols, such as bilobol or adipostatin A (bilobol is a substance related to anacardic acid from cashew nut shells and present in poison ivy and other Toxicodendron spp.) Individuals with a history of strong allergic reactions to poison ivy, mangoes, cashews and other alkylphenol-producing plants are more likely to experience an allergic reaction when consuming non-standardized ginkgo-containing preparations. The level of these allergens in standardized pharmaceutical preparations from Ginkgo biloba was restricted to 5 ppm by the Commission E of the former Federal German Health Authority. Overconsumption of seeds from Ginkgo biloba can deplete vitamin B6.
A 2021 umbrella review concluded that ginkgo biloba appears to be a beneficial and safe treatment for improving cognitive function and daily living activities in patients with Alzheimer’s disease. A 2018 systematic review and meta-analysis concluded that 22- to 24-week treatment with G. biloba extract EGb 761 significantly improved behavioral and psychological symptoms of dementia (excluding psychotic features) and reduced caregiver distress compared to placebo. A 2017 overview of systematic reviews concluded that G. biloba extract may modestly improve cognitive function and daily living in dementia when taken at doses over 200 mg/day for at least 22 weeks.
A 2016 systematic review concluded that significantly reduces tardive dyskinesia symptoms in schizophrenia patients and is generally safe.
It lacks United States Food and Drug Administration approval and lacks strong evidence to support its clinical use according to a 2023 review. The United States National Center for Complimentary and Integrative Health concludes that despite extensive research, ginkgo has not been conclusively shown to be effective for any health condition, including dementia, cognitive decline, or other issues for which it is often marketed.
The Committee on Herbal Medicinal Products (HMPC) in the European Union has concluded, based on clinical trials, that ginkgo leaf extract is effective and safe for improving cognitive function and quality of life in mild dementia, and it is licensed as a medicine in member states of the EU.
In professional sumo, wrestlers ranked in the two highest divisions ( and ) wear an elaborate chonmage called because it resembles the leaf of the ginkgo tree.
Ginkgo is an official tree of Seoul since 1971, designated by the Seoul Metropolitan Government.
==Gallery==
Classification
The relationship of ginkgo to other plant groups remains uncertain. It has been placed loosely in the divisions Spermatophyta and Pinophyta, but no consensus has been reached. Since its seeds are not protected by an ovary wall, it can morphologically be considered a gymnosperm. The apricot-like structures produced by female ginkgo trees are technically not , but are seeds that have a shell consisting of a soft and fleshy section (the sarcotesta), and a hard section (the sclerotesta). The sarcotesta has a strong smell that most people find unpleasant.
Approximate reconstructions by B. M. Begović Bego and Z. Zhou, 2010/2011. Source: B.M. Begović Bego, (2011). Nature's Miracle Ginkgo biloba , Book 1, Vols. 1–2, pp. 60–61.]]'']]
Phylogeny
Ginkgo biloba is a living fossil, with fossils recognisably related to modern ginkgo from the early Permian (Cisuralian), with likely oldest record being that of Trichopitys from the earliest Permian (Asselian) of France, over 290 million years old. The closest living relatives of the clade are the cycads, which share with the extant G. biloba the characteristic of motility sperm.
Rise and decline
Fossils attributable to the genus Ginkgo first appeared in the Middle Jurassic. The genus Ginkgo diversified and spread throughout Laurasia during the Jurassic and Early Cretaceous.
Limited number of species
It is doubtful whether the Northern Hemisphere fossil species of Ginkgo can be reliably distinguished. Given the slow pace of evolution and morphological similarity between members of the genus, there may have been only one or two species existing in the Northern Hemisphere through the entirety of the Cenozoic: present-day G. biloba (including G. adiantoides) and Ginkgo gardneri from the Paleocene of Scotland.
Adaptation to a single environment
Given the slow rate of evolution of the genus, Ginkgo possibly represents a pre-angiosperm strategy for survival in disturbed streamside environments. Ginkgo evolved in an era before flowering plants, when , , and Bennettitales dominated disturbed streamside environments, forming low, open, shrubby canopies. Ginkgo large seeds and habit of "bolting" – growing to a height of 10 meters before elongating its side branches – may be adaptations to such an environment.
Etymology
The older Chinese name for this plant is 銀果, meaning "silver fruit", pronounced yínguǒ in Mandarin or Ngan-gwo in Cantonese. The current commonly used names are 白果 (), meaning "white fruit", and 銀杏 (), meaning "silver apricot". The name 銀杏 was translated into Japanese as イチョウ () or ぎんなん () and into Korean language as 은행 ().
Distribution and habitat
Although Ginkgo biloba and other species of the genus were once widespread throughout the world, its habitat had shrunk by two million years ago.
Cultivation
Ginkgo has long been cultivated in China. It is common in the southern third of the country. Some planted trees at temples are believed to be over 1,500 years old. The first record of Europeans encountering it is in 1690 in Japanese garden, where the tree was seen by the German botanist Engelbert Kaempfer. Because of its status in Buddhism and Confucianism, the ginkgo has also been widely planted in Korea and in Japan since the 14th century; in both areas, some naturalization has occurred, with ginkgos seeding into natural forests. Ginkgo has been commonly cultivated in North America for over 200 years and in Europe for close to 300, but during that time, it has never become significantly naturalized.
Hiroshima
Extreme examples of the ginkgo's tenacity may be seen in Hiroshima, Japan, where six trees growing between from the 1945 atom bomb explosion were among the few living in the area to survive the blast. Although almost all other plants (and animals) in the area were killed, the ginkgos, though charred, survived and were soon healthy again, among other hibakujumoku (trees that survived the blast).
1000-year-old ginkgo at Tsurugaoka Hachimangū
At the Tsurugaoka Hachiman-gū's shrine in the city of Kamakura, Kanagawa Prefecture, Japan, an ancient ginkgo tree stands beside the stone entry staircase. According to legend, the tree has stood there since the founding of the shrine circa 1063. The tree is nicknamed kakure-ichō (hiding ginkgo), because of an Edo period legend in which shōgun Minamoto no Sanetomo was assassinated in 1219 by his nephew, Kugyō, who had hidden behind the tree to ambush the shōgun.
1,400-year-old ginkgo tree at Gu Guanyin
The grounds of the Buddhist temple at Gu Guanyin in the Zhongnan Mountains feature a ginkgo tree reputed to be 1,400 years old. The tree itself is a popular tourist attraction.
Toxicity
Since 2016, G. biloba extract is classified as a possible human carcinogen (group 2B) by the International Agency for Research on Cancer.
Uses
The wood of Ginkgo biloba is used to make furniture, chessboards, carving, and casks for making saké; the wood is fire-resistant and slow to decay.
Culinary
Despite the health risks in certain cases, the nut-like kernels of the seeds are esteemed in Asia, and are a traditional ingredient in Chinese cuisine. Ginkgo nuts are used in congee, and are often served at special occasions such as weddings and the Chinese New Year (as part of the vegetarian dish called Buddha's delight). Japanese cooks add ginkgo seeds (called ginnan) to dishes such as chawanmushi, and cooked seeds are often eaten along with other dishes. Grilled ginkgo nuts with salt are also a popular item at as a snack with beer and other Japanese food. In Korea, ginkgo nuts are stir-fried and eaten, or are used to garnish foods such as Sinseollo.
Clinical efficacy and safety
Although of G. biloba leaf are often marketed as cognitive enhancers, there is no evidence for effects on memory or attention in healthy people. Systematic reviews have shown there is no evidence for effectiveness of ginkgo in treating high blood pressure, menopause-related cognitive decline, tinnitus, post-stroke recovery, or altitude sickness.
Traditional medicine
Ginkgo has been used in traditional Chinese medicine since at least the 11th century CE. (2025). 9780300213829, Yale University Press. ISBN 9780300213829 Ginkgo seeds, leaves, and nuts have traditionally been used to treat various ailments, such as dementia, asthma, bronchitis, and kidney and bladder disorders. However, there is no conclusive evidence that ginkgo is useful for any of these conditions. (1997). 9789653500686, Akademon (Hebrew University of Jerusalem). ISBN 9789653500686, s.v. Ginkgo biloba
In culture
The ginkgo leaf is the symbol of the Urasenke school of Japanese tea ceremony. The tree is the official tree of the Japanese capital of Tokyo, and the symbol of Tokyo is a ginkgo leaf. Since 1948, the badge of Tokyo University has been two ginkgo leaves (designed by Shoichi Hoshino), which became the university logo in 2004 with a redesign. The logo of Osaka University has been a simplified ginkgo leaf since 1991 when designer Ikko Tanaka created it for the university's sixtieth anniversary.
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See also
External links
Categories: Biloba, Cyp3a4 Inhibitors, Dioecious Plants, Edible Plants, Endangered Plants, Endemic Flora Of China, Extant Ypresian First Appearances, Flora Of Zhejiang, Garden Plants Of Asia, Medicinal Plants Of Asia, Plants Used In Bonsai, Plants Used In Traditional Chinese Medicine, Taxa Named By Carl Linnaeus, Trees Of China, Plants Described In 1771, Trees
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