Thegarden strawberry (or simplystrawberry;Fragaria × ananassa)[1] is a widely grownhybrid cultivated worldwide for itsfruit. The genusFragaria, the strawberries, is in the rose family,Rosaceae. The fruit is appreciated for its aroma, bright red colour, juicy texture, and sweetness. It is eaten either fresh or in prepared foods such asjam,ice cream, andchocolates. Artificial strawberry flavourings and aromas are widely used in commercial products. Botanically, the strawberry is not aberry, but anaggregateaccessory fruit. Each apparent 'seed' on the outside of the strawberry is actually anachene, a botanical fruit with a seed inside it.
The garden strawberry was first bred inBrittany, France, in the 1750s via a cross ofF. virginiana from eastern North America andF. chiloensis, which was brought from Chile byAmédée-François Frézier in 1714.Cultivars ofF. ×ananassa have replaced the woodland strawberryF. vesca in commercial production. In 2022, world production of strawberries exceeded nine million tons, led by China with 35% of the total.
Strawberries have appeared in literature and art from Roman times;Virgil wrote about the snake lurking beneath the strawberry, an image reinterpreted by later writers includingShakespeare. Strawberries appear in Italian, Flemish, and German paintings, includingHieronymus Bosch'sThe Garden of Earthly Delights. It has been understood to symbolise the ephemerality of earthly joys or the benefit that blessed souls get from religion, or to allegorise death and resurrection. By the late 20th century, its meaning had shifted: it symbolised female sexuality.
Evolution
History and taxonomy
In Europe, until the 17th century cultivated plants were obtained by transplanting strawberries from the forests; the plants werepropagated asexually by pegging down therunners, allowing them to root, and then separating the new plants.[2]F. virginiana, the Virginia strawberry, was brought to Europe from eastern North America;F. chiloensis, the Chilean strawberry, was brought from Chile byAmédée-François Frézier in 1714. At first introduction to Europe, the Chilean strawberry plants grew vigorously, but produced no fruit. French gardeners inBrittany in the 1750s noticed that the Chilean plants bore only female flowers. They planted the wild woodland strawberryF. vesca among the Chilean plants to provide pollen; the Chilean strawberry plants then bore abundant fruits.[3]
In 1759, Philip Miller recorded the 'pine strawberry' (F. ananassa) inChelsea, England.[3] In the gardens of thePalace of Versailles, France,Antoine Nicolas Duchesne found in 1766 thatF. ananassa was a hybrid of the recently arrivedF. chiloensis andF. virginiana.[1] In 1806, Michael Keens ofIsleworth, England selected the Keens Imperial cultivar from many hybrids,[4] winning theRoyal Horticultural Society's Silver Cup.[3] Both the names 'pine' and 'ananassa' meant "pineapple", for the fruit's flavour.[4] Modern strawberries and both parent species areoctoploid (8N, meaning they have 8 sets of 7chromosomes).[5] Thegenome sequence of the garden strawberry was published in 2019.[6]
Hybridisation and polyploidy in strawberries. Garden strawberries areoctoploid (8N), like both parents, the Virginia and Chilean strawberries.
Further breeding in the following centuries produced varieties with a longer cropping season and more fruit.[3] During theGreen Revolution of the 1950s,agronomists used selective breeding to expandphenotypic diversity of the garden strawberry. Adoption of perpetual flowering hybrids not sensitive to changes inphotoperiod gave higher yields and enabled production in California to expand.[1]
Fragaria vesca, a wild woodland strawberry, was cultivated until the 17th century.
Botanical structure of a strawberry, compared to a peapod. The strawberry is a swollenreceptacle, covered with many smallachenes, the botanical fruits.[8]
In culinary terms, a strawberry is an edible fruit. From a botanical point of view, it is not aberry but anaggregateaccessory fruit, because the fleshy part is derived from thereceptacle. Each apparent seed on the outside of the strawberry is actually anachene, a botanical fruit with a seed inside it.[8]
Furaneol contributes to the fragrance of strawberries.
Sweetness, fragrance and complex flavour are important attributes of strawberries.[16] Inplant breeding and farming, emphasis is placed on sugars, acids, andvolatile compounds, which improve the taste and fragrance of the ripe fruit.[17]Esters,terpenes, andfurans are the chemical compounds having the strongest relationships to strawberry flavour, sweetness and fragrance, with a total of 31 out of some 360 volatile compounds significantly correlated to desirable flavour and fragrance.[17][18][19] In breeding strawberries for the commercial market in the United States, the volatile compoundsmethyl anthranilate andgamma-decalactone, prominent in aromatic wild strawberries, are especially desired for their "sweet and fruity" aroma characteristics.[18][19] As strawberry flavour andfragrance appeal to consumers,[18][19][20] they are used widely in manufacturing, including foods, beverages, perfumes and cosmetics.[21][22]
Allergy
Some people experience ananaphylactoid reaction to eating strawberries.[23] The most common form of this reaction isoral allergy syndrome, but symptoms may also mimichay fever or includedermatitis orhives, and, in severe cases, may cause breathing problems.[24]Proteomic studies indicate that the allergen may be tied to a protein for the red anthocyanin biosynthesis expressed in strawberry ripening, named Fra a1 (Fragaria allergen1). White-fruited strawberrycultivars, lacking Fra a1, may be an option for people allergic to strawberries.[25] They ripen but remain pale, appearing like immature berries. A virtually allergen-free cultivar named 'Sofar' is available.[26][27]
Strawberries are often grouped according to their flowering habit.[28][29] Traditionally in the Northern Hemisphere, this has consisted of a division between "June-bearing" strawberries, which bear their fruit in the early summer and "everbearing" strawberries, which often bear several crops of fruit throughout the season.[29] One plant throughout a season may produce 50 to 60 times or roughly once every three days.[30] Strawberries occur in three basic flowering habits: short-day, long-day, and day-neutral. These describe the day-length sensitivity of the plant and the type ofphotoperiod that induces flower formation. Day-neutral cultivars produce flowers regardless of the photoperiod.[31] Strawberry cultivars vary widely in size, colour, flavour, shape, degree of fertility, season of ripening, liability to disease and constitution of plant.[28]
In 2022, world production of strawberries was 9.6milliontonnes, led by China with 35 percent of the total and the United States andTurkey as other significant producers.[32] Due to the relatively fragile nature of the strawberry, approximately 35 percent of the $2.2billion United States crop was spoiled in 2020. AnIdaho company announced plans to launch more durable gene-edited strawberries. In the U.S., as of 2021[update], it cost growers around $35,000 per acre to plant and $35,000 per acre to harvest strawberries.[33]
For commercial production, plants can be propagated from bare root plants or plugs. One method of cultivation uses annualplasticulture;[34] another is a perennial system of matted rows or mounds which has been used in cold growing regions for many years.[35] In some areas, greenhouses are used; in principle they could provide strawberries during the off season for field crops.[36]
In the plasticulture system, raised beds are covered with plastic to prevent weed growth and erosion. Plants are planted through holes punched in this covering. Irrigation tubing can be run underneath if necessary.[34][37]
Strawberries may be propagated by seed.[40] Strawberries can be grown indoors in pots.[41] Strawberries will not grow indoors in winter though an experiment using a combination of blue and redLED lamps shows that this could be achieved in principle.[42] InFlorida, winter is the natural growing season and harvesting begins in mid-November.[30]
Strawberry field in North Rhine-Westphalia, Germany
A field using the plasticulture method
Manuring and harvesting
Strawberries are usually picked and placed in shallow boxes in the field.
Nitrogen fertiliser is often needed at the beginning of every planting year. There are normally adequate levels of phosphorus and potash when fields have been fertilised for other crops in preceding years. To provide more organic matter, a cover crop of wheat or rye can be planted in the year before planting the strawberries. Strawberries prefer a somewhat acidic pH from 5.5 to 6.5, so lime is usually not required.[43]
To achieve top quality, berries are harvested at least every other day. The berries are picked with the caps and half the stem still attached. Strawberries need to remain on the plant until fully ripe, because they do not continue to ripen after being picked.[44] The harvesting and cleaning process has not changed substantially over time. As they are delicate, strawberries are still often harvested by hand and packed in the field.[45]
TheNPR1 gene fromArabidopsis thaliana,AtNPR1, confersA. thaliana'sbroad-spectrum resistance whentransexpressed inF. ananassa. This includes resistance to anthracnose, powdery mildew, and angular leaf spot.[59]
A 1997 study found that manywound volatiles were effective against gray mold (B. cinerea). BothTribute andChandler varieties benefited from the treatments, although the effects vary widely with substance and variety. Strawberry plantsmetabolise these volatiles, more rapidly than do eitherblackberry orgrape.[60]
Strawberries were eaten fresh with cream in the time ofThomas Wolsey in the court of KingHenry VIII.[61] Strawberries can be frozen or made into jam orpreserves,[62] as well as dried and used in prepared foods, such as cereal bars.[63] In the United Kingdom, strawberries and cream is a popular dessert at theWimbledon tennis tournament.[61] Desserts using strawberries includepavlova,[64]fraisier,[65][66] and strawberryshortcake.[67]
The Roman poetOvid wrote that in the past Golden Age, people had lived on wild fruits such as mountain strawberries.[68]Virgil wrote in hisEclogues that "Ye who cull flowers and low-growing strawberries, / Away from here lads; a chill snake lurks in the grass", and his imagery was taken up by medieval andearly modern writers, the snake beneath the strawberry standing for dangerous literature, or beautiful but unfaithful women, or eventually any risky pleasure. In this vein,Shakespeare's KingRichard III asks for a dish of strawberries while feigning friendship to his enemy; while inOthello,Iago showsDesdemona's handkerchief "spotted with strawberries", implying she has been unfaithful and hinting at Iago's own devious plans.[69]
The strawberry is found in Italian, Flemish, and German art, and in English miniatures.[69] In medieval depictions, the strawberry often appears in theVirgin Mary's garden, while in theMadonna of the Strawberries, she is seated on a strawberry bed and garlanded with strawberry leaves.[69]
In the work of the late medieval painterHieronymus Bosch, strawberries feature inThe Garden of Earthly Delights amongst "frolicking nude figures".[68] Fray Jose de Siguenza described the painting as embodying the strawberry as a symbol of theephemerality of earthly joys.[68] More recently, scholars have seen the symbolism entirely differently: Clément Wertheim-Aymes believed it meant the blessed souls' benefit from religion; Pater Gerlach supposed it meant spiritual love; and Laurinda Dixon asserted it was part of anallegory of death and resurrection.[68] By the late 20th century, the strawberry (and theraspberry) had become "traditional symbols of the mouth and female sexuality".[70]
^Vrhovsek, U.; Guella, G.; Gasperotti, M.; Pojer, E.; Zancato, M.; Mattivi, F. (2012). "Clarifying the Identity of the Main Ellagitannin in the Fruit of the Strawberry,Fragaria vesca andFragaria ananassa Duch".Journal of Agricultural and Food Chemistry.60 (10):2507–2516.Bibcode:2012JAFC...60.2507V.doi:10.1021/jf2052256.PMID22339338.
^Aaby, K.; Skrede, G.; Wrolstad, R.E. (2005). "Phenolic composition and antioxidant activities in flesh and achenes of strawberries (Fragaria ananassa)".Journal of Agricultural and Food Chemistry.53 (10):4032–4040.Bibcode:2005JAFC...53.4032A.doi:10.1021/jf048001o.PMID15884835.
^Fossen, Torgils; Rayyan, Saleh; Andersen, Øyvind M. (2004). "Dimeric anthocyanins from strawberry (Fragaria ananassa) consisting of pelargonidin 3-glucoside covalently linked to four flavan-3-ols".Phytochemistry.65 (10):1421–1428.Bibcode:2004PChem..65.1421F.doi:10.1016/j.phytochem.2004.05.003.PMID15231416.
Silva, Katchen Julliany P.; Brunings, Asha; Peres, Natalia A.; Mou, Zhonglin; Folta, Kevin M. (27 March 2015). "The Arabidopsis NPR1 gene confers broad-spectrum disease resistance in strawberry".Transgenic Research.24 (4). Springer Science and Business Media LLC:693–704.doi:10.1007/s11248-015-9869-5.PMID25812515.
^Giampieri, Francesca; Alvarez-Suarez, Josè M.; Mazzoni, Luca; Romandini, Stefania; Bompadre, Stefano; et al. (2013). "The potential impact of strawberry on human health".Natural Product Research.27 (4–5). Informa UK:448–455.doi:10.1080/14786419.2012.706294.PMID22788743.
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