Cucurbita (Latin for 'gourd')[2][3] is agenus ofherbaceousfruits in the gourdfamily,Cucurbitaceae (also known ascucurbits orcucurbi), native to theAndes andMesoamerica. Five edible species are grown and consumed for their flesh and seeds. They are variously known assquash,pumpkin, or gourd, depending on species,variety, and local parlance.[a] Other kinds of gourd, also called bottle-gourds, are native to Africa and belong to the genusLagenaria, which is in the same family and subfamily asCucurbita, but in a differenttribe; their young fruits are eaten much like those of theCucurbita species.
MostCucurbita species are herbaceous vines that grow several meters in length and havetendrils, but non-vining "bush" cultivars ofC. pepo andC. maxima have also been developed. The yellow or orange flowers on aCucurbita plant are of two types:female and male. The female flowers produce the fruit and the male flowers producepollen. Many North and Central American species are visited by specialistbeepollinators, but other insects with more general feeding habits, such ashoney bees, also visit.
The fruits of the genusCucurbita are good sources ofnutrients, such asvitamin A andvitamin C, among other nutrients according to species. The fruits have many culinary uses includingpumpkin pie,biscuits,bread,desserts,puddings,beverages, andsoups; they are nowcultivated worldwide. Although botanical fruits,Cucurbita gourds such as squash are typically cooked and eaten asvegetables. Pumpkins see more varied use, and are eaten both as vegetables and as desserts such as pumpkin pie.
Cucurbita species fall into two main groups. The first group consists ofannual or short-livedperennial vines which aremesophytic, meaning they require a more or less continuous water supply. The second group areperennials growing in arid zones which arexerophytic, meaning they tolerate dry conditions. CultivatedCucurbita species were derived from the first group. Growing 5 to 15 meters (15 to 50 feet) in height or length, the plant stem producestendrils to help it climb adjacent plants and structures or extend along the ground. Most species do not readily root from the nodes; a notable exception isC. ficifolia, and the four other cultivated mesophytes do this to a lesser extent. The vine of the perennialCucurbita can become semiwoody if left to grow. There is wide variation in size, shape, and color amongCucurbita fruits, and even within a single species.C. ficifolia is an exception, being highly uniform in appearance.[5] Themorphological variation in the speciesC. pepo[6] andC. maxima[7] is so vast that its varioussubspecies and cultivars have been misidentified as totally separate species.[6]
The typical cultivatedCucurbita species has five-lobed orpalmately divided leaves with longpetioles, with the leaves alternately arranged on the stem. The stems in some species are angular. All of the above-ground parts may be hairy with various types oftrichomes, which are often hardened and sharp. Spring-like tendrils grow from eachnode and are branching in some species.C. argyrosperma has ovate-cordate (egg-shaped to heart-shaped) leaves. The shape ofC. pepo leaves varies widely.C. moschata plants can have light or densepubescence.C. ficifolia leaves are slightly angular and have light pubescence. The leaves of all four of these species may or may not have white spots.[8]
The species aremonoecious, with unisexual male (staminate) and female (pistillate) flowers on a single plant and these grow singly, appearing from theleaf axils. Flowers have five fused yellow to orange petals (thecorolla) and a green bell-shapedcalyx. Male flowers in Cucurbitaceae generally have five stamens, but inCucurbita there are only three, and theiranthers are joined so that there appears to be one.[9][10] Female flowers have thickpedicels, and aninferior ovary with 3–5stigmas that each have two lobes.[8][11] The female flowers ofC. argyrosperma andC. ficifolia have larger corollas than the male flowers.[8] Female flowers ofC. pepo have a small calyx, but the calyx ofC. moschata male flowers is comparatively short.[8]
Cucurbita fruits are large and fleshy.[9] Botanists classify theCucurbita fruit as apepo, which is a special type ofberry derived from an inferior ovary, with a thick outer wall or rind withhypanthium tissue forming anexocarp around the ovary, and a fleshy interior composed ofmesocarp andendocarp. The term "pepo" is used primarily for Cucurbitaceae fruits, where this fruit type is common, but the fruits ofPassiflora andCarica are sometimes also pepos.[12][13] The seeds, which are attached to the ovary wall (parietal placentation) and not to the center, are large and fairly flat with a large embryo that consists almost entirely of twocotyledons.[11] Fruit size varies considerably: wild fruit specimens can be as small as 4 centimeters (1+1⁄2 inches) and some domesticated specimens can weigh well over 300 kilograms (660 pounds).[8] The current world record was set in 2014 by Beni Meier ofSwitzerland with a 1,054 kg (2,323.7 lb) pumpkin.[14]
Male flower, part of the perianth removed, arrows indicating nectar pores
When there is more pollen applied to the stigma, more seeds are produced in the fruits and the fruits are larger with greater likelihood of maturation,[18] an effect calledxenia. Competitively grown specimens are therefore often hand-pollinated to maximize the number of seeds in the fruit.[19][20]Seedlessness is known to occur in certain cultivars ofC. pepo.[21][22]
Critical factors in flowering and fruit set are physiological, having to do with the age of the plant and whether it already has developing fruit.[23] Theplant hormonesethylene andauxin are key in fruit set and development.[24] Ethylene promotes the production of female flowers. When a plant already has a fruit developing, subsequent female flowers on the plant are less likely to mature, a phenomenon called "first-fruit dominance",[23] and male flowers are more frequent, an effect that appears due to reduced natural ethylene production within the plant stem.[25]Ethephon, a plant growth regulator product that is converted to ethylene after metabolism by the plant, can be used to increase fruit and seed production.[19][26] AlthoughCucurbita species can generally produce healthy fruit after pollination from the same plant,inbreeding depression can significantly reduce seed number and fruit size.[27]
The plant hormonegibberellin, produced in the stamens, is essential for the development of all parts of the male flowers. The development of female flowers is not yet understood.[28] Gibberellin is also involved in other developmental processes of plants, such as seed and stem growth.[29]
Seeds with maximumgermination potential develop (inC. moschata) by 45 days afteranthesis, and seed weight reaches its maximum 70 days after anthesis.[30] Some varieties ofC. pepo germinate best with eight hours of sunlight daily and a planting depth of12 millimeters (1⁄2 in). Seeds planted deeper than 125 millimeters (5 in) are not likely to germinate.[31] InC. foetidissima, a weedy species, plants younger than 19 days old are not able to sprout from the roots after removing the shoots. In a seed batch with 90 percent germination rate, over 90 percent of the plants had sprouted after 29 days from planting.[32]
Experiments have shown that when more pollen is applied to the stigma, as well as the fruit containing more seeds and being larger (the xenia effect mentioned above), the germination of the seeds is also faster and more likely, and the seedlings are larger.[18] Various combinations of mineral nutrients and light have a significant effect during the various stages of plant growth. These effects vary significantly between the different species ofCucurbita. A type of stored phosphorus calledphytate forms in seed tissues as spherical crystalline intrusions in protein bodies calledgloboids. Along with other nutrients, phytate is used completely during seedling growth.[33]Heavy metal contamination, includingcadmium, has a significant negative impact on plant growth.[34]Cucurbita plants grown in the spring tend to grow larger than those grown in the autumn.[35]
Cucurbita was formally described in a way that meets the requirements of modernbotanical nomenclature byLinnaeus in hisGenera Plantarum,[36] the fifth edition of 1754 in conjunction with the 1753 first edition ofSpecies Plantarum.[37]Cucurbita pepo is thetype species of the genus.[37][38] Linnaeus initially included the speciesC. pepo,C. verrucosa andC. melopepo (both now included inC. pepo), as well asC. citrullus (watermelon, nowCitrullus lanatus) andC. lagenaria (nowLagenaria siceraria) (both are notCucurbita but are in the family Cucurbitaceae.[39]
TheCucurbita digitata,C. foetidissima,C. galeotti, andC. pedatifoliaspecies groups arexerophytes, arid zone perennials with storage roots; the remainder, including the five domesticated species, are all mesophytic annuals or short-life perennials with no storage roots.[5][40] The five domesticated species are mostly isolated from each other bysterility barriers and have different physiological characteristics.[40] Somecross pollinations can occur:C. pepo withC. argyrosperma andC. moschata; andC. maxima withC. moschata. Cross pollination does occur readily within the family Cucurbitaceae.[41] The buffalo gourd (C. foetidissima) has been used as an intermediary, as it can be crossed with all the commonCucurbita.[11]
An assortment of fruits ofC. maxima andC. pepo
Varioustaxonomic treatments have been proposed forCucurbita, ranging from 13 to 30 species.[2] In 1990,Cucurbita expert Michael Nee classified them into the following oft-cited 13 species groups (27 species total), listed by group and alphabetically, with geographic origin:[5][42][43][44]
The taxonomy by Nee closely matches the species groupings reported in a pair of studies by a botanical team led by Rhodes and Bemis in 1968 and 1970 based on statistical groupings of severalphenotypic traits of 21 species. Seeds for studying additional species members were not available. Sixteen of the 21 species were grouped into five clusters with the remaining five being classified separately:[15][47]
C. digitata,C. palmata,C. californica,C. cylindrata,C. cordata
C. martinezii,C. okeechobeensis,C. lundelliana
C. sororia,C. gracilior,C. palmeri;C. argyrosperma (reported asC. mixta) was considered close to the three previous species
C. maxima,C. andreana
C. pepo,C. texana
C. moschata,C. ficifolia,C. pedatifolia,C. foetidissima, andC. ecuadorensis were placed in their own separate species groups as they were not considered significantly close to any of the other species studied.
The fullphylogeny of this genus is unknown, and research was ongoing in 2014.[48][49] The followingcladogram ofCucurbita phylogeny is based upon a 2002 study ofmitochondrial DNA by Sanjur and colleagues.[50]
A festival-winning pumpkin in 2009 weighing 742 kilograms (1,636 lb)
The ancestral species of the genusCucurbita were present in the Americas before thearrival of humans,[51][52] and are native to theAmericas. The likely center of origin is southern Mexico, spreading south through what is now known as Mesoamerica, into South America, and north to what is now the southwestern United States.[51] Evolutionarily speaking, the genus is relatively recent in origin, dating back to theHolocene, whereas the family Cucurbitaceae, represented inBryonia-like seeds, dates to thePaleocene.[53] Recent genomic studies support the idea that theCucurbita genus underwent a whole-genome duplication event, increasing the number of chromosomes and accelerating the rate at which their genomes evolve relative to other cucurbits.[54][55][56] No species within the genus is entirely genetically isolated.C. moschata can intercross with allCucurbita species, though the hybrid offspring may not be fertile unless they becomepolyploid.[40]
Evidence of domestication of Cucurbita goes back over 8,000 years from the southernmost parts of Canada down to Argentina and Chile. Centers of domestication stretch from the Mississippi River watershed and Texas down through Mexico and Central America to northern and western South America.[5] Of the 27 species that Nee delineates, five are domesticated. Four of these,C. argyrosperma,C. ficifolia,C. moschata, andC. pepo, originated and were domesticated inMesoamerica; the fifth,C. maxima, originated and was domesticated in South America.[8]
WithinC. pepo, the pumpkins, thescallops, and possibly the crooknecks are ancient and were domesticated at different times and places. The domesticated forms ofC. pepo have larger fruits than non-domesticated forms and seeds that are larger but fewer in number.[57] In a 1989 study on the origins and development ofC. pepo, botanist Harry Paris suggested that the original wild specimen had a small round fruit and that the modern pumpkin is its direct descendant. He suggested that the crookneck, ornamental gourd, and scallop are early variants and that the acorn squash is a cross between the scallop and the pumpkin.[57]
C. moschata 'Butternut'
C. argyrosperma is not as widespread as the other species. The wild formC. a. subsp.sororia is found fromMexico toNicaragua, and cultivated forms are used in a somewhat wider area stretching from Panama to the southeastern United States.[8] It was probably bred for its seeds, which are large and high inoil andprotein, but its flesh is of poorer quality than that ofC. moschata andC. pepo. It is grown in a wide altitudinal range: from sea level to as high as 1,800 meters (5,900 ft) in dry areas, usually with the use of irrigation, or in areas with a defined rainy season, where seeds are sown in May and June.[8]
C. ficifolia andC. moschata were originally thought to be Asiatic in origin, but this has been disproven. The origin ofC. ficifolia is Latin America, most likely southern Mexico, Central America, or the Andes. It grows at elevations ranging from 1,000 to 3,000 meters (3,300 to 9,800 ft) in areas with heavy rainfall. It does not hybridize well with other cultivated species as it has significantly different enzymes and chromosomes.[8]
C. maxima originated in South America over 4,000 years ago,[50] probably in Argentina and Uruguay. The plants are sensitive tofrost, and they prefer both bright sunlight and soil with a pH of 6.0 to 7.0.[58]C. maxima did not start to spread into North America until after the arrival of Columbus. Varieties were in use by native peoples of the United States by the 16th century.[5] Types ofC. maxima includetriloba,[59]zapallito,[60]zipinka,[61] Banana, Delicious, Hubbard, Marrow (C. maxima Marrow), Show, and Turban.[62]
Fruit of the 'Tromboncino' cultivar of the Crookneck (C. moschata)Group are eaten either when very young, or as mature winter squash.
C. moschata is native to Latin America, but the precise location of origin is uncertain.[63] It has been present in Mexico, Belize, Guatemala, and Peru for 4,000–6,000 years and has spread to Bolivia, Ecuador, Panama, Puerto Rico, and Venezuela. This species is closely related toC. argyrosperma. A variety known as the Seminole Pumpkin has been cultivated in Florida since before the arrival of Columbus. Its leaves are 20 to 30 centimeters (8 to 12 in) wide. It generally grows at low elevations in hot climates with heavy rainfall, but some varieties have been found above 2,200 meters (7,200 ft).[8] Groups ofC. moschata include Cheese, Crookneck (C. moschata), and Bell.[62]
C. pepo is one of the oldest, if not the oldest, domesticated species with the oldest known locations beingOaxaca, Mexico, 8,000–10,000 years ago, andOcampo, Tamaulipas, Mexico, about 7,000 years ago. It is known to have appeared inMissouri, United States, at least 4,000 years ago.[5][8][64][65] Debates about the origin ofC. pepo have been on-going since at least 1857.[66] There have traditionally been two opposing theories about its origin: 1) that it is a direct descendant ofC. texana and 2) thatC. texana is merelyferalC. pepo.[5] A more recent theory by botanist Thomas Andres in 1987 is that descendants ofC. fraterna hybridized withC. texana,[67] resulting in two distinct domestication events in two different areas: one in Mexico and one in the eastern United States, withC. fraterna andC. texana, respectively, as the ancestral species.[8][50][67][68]C. pepo may have appeared in the Old World before moving from Mexico into South America.[8] It is found from sea level to slightly above 2,000 meters (6,600 ft). Leaves have 3–5 lobes and are 20–35 centimeters (8–14 in) wide. All the subspecies, varieties, andcultivars areinterfertile.[6] In 1986 Paris proposed a revised taxonomy of the edible cultivatedC. pepo based primarily on the shape of the fruit, with eight groups.[57][69] All but a fewC. pepo cultivars can be included in these groups.[8][69][70][71] There is one non-edible cultivated variety:C. pepo var.ovifera.[72]
A classification of cultivatedC. pepo varieties based on Paris' eight groups and the one non-edible variety
Winter squash, both a shrubby and creeping plant, obovoid or conical shape, pointed at the apex and with longitudinal grooves, thus resembling a spinning top,[69] ex:Acorn squash[8][70][71]
Cocozzelle
C. pepo var.Ionga
Summer squash, long round slender fruit that is slightly bulbous at the apex,[69] similar tofastigata, ex: Cocozelle von tripolis[8][70][71]
Crookneck
C. pepo var.torticollia (alsotorticollis)
Summer squash, shrubby plant, with yellow, golden, or white fruit which is long and curved at the end and generally has a verrucose (wart-covered) rind,[69] ex:Crookneck squash[8][70][71]
Pumpkin
C. pepo var.pepo
Winter squash, creeping plant, round, oblate, or oval shape and round or flat on the ends,[69] ex:Pumpkin;[8][70][71] includesC. pepo subsp.pepo var.styriaca, used for Styrianpumpkin seed oil[73]
Scallop
C. pepo var.clypeata; calledC. melopepo byLinnaeus[6]
Summer squash, prefers half-shrubby habitat, flattened or slightly discoidal shape, with undulations or equatorial edges,[69] ex:Pattypan squash[8][70][71]
Straightneck
C. pepo var.recticollis
Summer squash, shrubby plant, with yellow or golden fruit and verrucose rind, similar to var.torticollia but a stem end that narrows,[69] ex:Straightneck squash[8][70][71]
Vegetable marrow
C. pepo var.fastigata
Summer and winter squashes, creeper traits and a semi-shrub, cream to dark green color, short round fruit with a slightly broad apex,[69] ex:Spaghetti squash (a winter variety)[8][70][71]
Zucchini/Courgette
C. pepo var.cylindrica
Summer squash, presently the most common group of cultivars, origin is recent (19th century), semi-shrubby, cylindrical fruit with a mostly consistent diameter,[69] similar tofastigata, ex:Zucchini[8][70][71]
Ornamental gourds
C. pepo var.ovifera
Non-edible,[72] field squash closely related toC. texana, vine habitat, thin stems, small leaves, three sub-groups:C. pepo var.ovifera (egg-shaped, pear-shaped),C. pepo var.aurantia (orange color), andC. pepo var.verrucosa (round warty gourds), ornamental gourds found in Texas and called var.texana and ornamental gourds found outside of Texas (Illinois, Missouri, Arkansas, Oklahoma, and Louisiana) are called var.ozarkana[64]
The genus was part of the culture of almost every native peoples group from southern South America to southern Canada.[52] Modern-day cultivatedCucurbita are not found in the wild.[5] Genetic studies of themitochondrial genenad1 show there were at least six independent domestication events ofCucurbita separating domestic species from their wild ancestors.[50] Species native to North America includeC. digitata (calabazilla),[88] andC. foetidissima (buffalo gourd),[89]C. palmata (coyote melon), andC. pepo.[5] Some species, such asC. digitata andC. ficifolia, are referred to asgourds. Gourds, also called bottle-gourds, which are used as utensils or vessels, belong to the genusLagenaria and are native to Africa.Lagenaria are in the same family and subfamily asCucurbita but in a differenttribe.[90]
The earliest known evidence of the domestication ofCucurbita dates back at least 8,000 years ago, predating the domestication of other crops such asmaize andbeans in the region by about 4,000 years.[5][64][65][91] This evidence was found in theGuilá Naquitz cave in Oaxaca, Mexico, during a series of excavations in the 1960s and 1970s, possibly beginning in 1959.[92][93] Solid evidence of domesticatedC. pepo was found in the Guilá Naquitz cave in the form of increasingrind thickness and largerpeduncles in the newerstratification layers of the cave. By c. 8,000 yearsBP theC. pepo peduncles found are consistently more than10 millimeters (3⁄8 in) thick. WildCucurbita peduncles are always below this 10 mm barrier. Changes in fruit shape and color indicate that intentional breeding ofC. pepo had occurred by no later than 8,000 years BP.[11][94][95] During the same time frame, average rind thickness increased from0.84–1.15 millimeters (1⁄32–3⁄64 in).[96] Recent genomic studies suggest thatCucurbita argyrosperma was domesticated in Mexico, in the region that is currently known as the state ofJalisco.[97]
*includes pumpkins Source:FAOSTAT of the United Nations[102]
In 2021, world production of squashes (including gourds and pumpkins) was 23.4 million tonnes, led by China with 32% of the total (table). Ukraine, Russia, and the United States were secondary producers.
Cucurbitin is anamino acid and acarboxypyrrolidine that is found in rawCucurbita seeds.[103][104] It retards the development of parasiticflukes when administered to infected host mice, although the effect is seen only if administration begins immediately after infection.[105]
Cucurbitacin is aplant steroid present in wildCucurbita and in each member of the familyCucurbitaceae. Poisonous to mammals,[79] it is found in quantities sufficient to discourage herbivores. It makes wildCucurbita and most ornamental gourds, with the exception of an occasionalC. fraterna andC. sororia, bitter to taste.[2][67][108] Ingesting too much cucurbitacin can cause stomach cramps, diarrhea and even collapse.[109] This bitterness is especially prevalent in wildCucurbita; in parts of Mexico, the flesh of the fruits is rubbed on a woman's breast to wean children.[110] While the process of domestication has largely removed the bitterness from cultivated varieties,[2] there are occasional reports of cucurbitacin causing illness in humans.[2] Cucurbitacin is also used as a lure in insect traps.[108]
As an example ofCucurbita, raw summer squash is 94% water, 3%carbohydrates, and 1%protein, with negligiblefat content (table). In a 100-gram reference serving, raw squash supplies 69 kilojoules (16 kcal) offood energy and is rich invitamin C (20% of theDaily Value, DV), moderate invitamin B6 andriboflavin (12–17% DV), but otherwise devoid of appreciable nutrient content (table), although the nutrient content of differentCurcubita species may vary somewhat.[113]
The family Cucurbitaceae has many species used as human food.[8]Cucurbita species are some of the most important of those, with various species being prepared and eaten in many ways. Although the stems and skins tend to be more bitter than the flesh,[109][117] the fruits andseeds of cultivated varieties are usually quite edible and need little or no preparation. Cross-pollination with toxic types can cause bitterness in plants of the next generation, and these should not be eaten. The flowers and young leaves and shoot tips can also be consumed.[118] The seeds and fruits of most varieties can be stored for long periods of time,[5] particularly the sweet-tasting winter varieties with their thick, inedible skins.[119] Summer squash have a thin, edible skin. The seeds of both types can be roasted, eaten raw, made intopumpkin seed oil,[73] ground into a flour or meal,[120] or otherwise prepared. Squashes are primarily grown for the fresh food market.[121]
Pumpkin custard made from kabocha, a cultivated variant ofC. maxima
Long before European contact,Cucurbita had been a major food source for the native peoples of the Americas. The species became an important food for European settlers, including thePilgrims, who even featured it at the firstThanksgiving.[11] Commercially produced pumpkin commonly used inpumpkin pie is most often varieties ofC. moschata;Libby's, by far the largest producer of processed pumpkin, uses a proprietary strain of the Dickinson pumpkin variety ofC. moschata for its canned pumpkin.[122] Other foods that can be made using members of this genus include biscuits,bread, cheesecake, desserts, donuts,granola, ice cream, lasagna dishes, pancakes, pudding, pumpkin butter,[123] salads, soups, and stuffing.[124]Squash soup is a dish in African cuisine.[125] Thexerophytic species are proving useful in the search for nutritious foods that grow well in arid regions.[126]C. ficifolia is used to make soft and mildly alcoholic drinks.[8]
In India, squashes (ghiya) are cooked with seafood such as prawns.[127] In France, marrows (courges) are traditionally served as agratin, sieved and cooked with butter, milk, and egg, and flavored with salt, pepper, and nutmeg,[128] and as soups. In Italy, zucchini and larger squashes are served in a variety of regional dishes, such ascocuzze alla puviredda cooked with olive oil, salt and herbs fromApulia; astorta di zucca fromLiguria, ortorta di zucca e riso fromEmilia-Romagna, the squashes being made into a pie filling with butter,ricotta,Parmesan, egg, and milk; and as a sauce for pasta in dishes likespaghetti alle zucchine from Sicily.[129] In Japan, squashes such as smallC. moschata pumpkins (kabocha) are eaten boiled with sesame sauce, fried as atempura dish, or made into balls withsweet potato andJapanese mountain yam.[130]
Along with maize and beans, squash has been depicted in the art work of the native peoples of the Americas for at least 2,000 years.[131][132] For example, cucurbits are often represented inMoche ceramics.[131][133]
Though native to the western hemisphere,Cucurbita began to spread to other parts of the world afterChristopher Columbus's arrival in the New World in 1492.[134][135] Until recently, the earliest known depictions of this genus in Europe was ofCucurbita pepo inDe Historia Stirpium Commentarii Insignes in 1542 by the German botanistLeonhart Fuchs, but in 1992, two paintings, one ofC. pepo and one ofC. maxima, painted between 1515 and 1518, were identified infestoons atVilla Farnesina in Rome.[136] Also, in 2001 depictions of this genus were identified inGrandes Heures of Anne of Brittany (Les Grandes Heures d'Anne de Bretagne), a French devotional book, anilluminated manuscript created between 1503 and 1508. This book contains an illustration known asQuegourdes de turquie, which was identified by cucurbit specialists asC. pepo subsp.texana in 2006.[137]
In 1952, Stanley Smith Master, using the pen name Edrich Siebert, wrote "The Marrow Song (Oh what a beauty!)" to a tune in6 8 time. It became a popular hit in Australia in 1973,[138] and was revived bythe Wurzels in Britain on their 2003 albumCutler of the West.[139][140]John Greenleaf Whittier wrote a poem entitledThe Pumpkin in 1850.[141] "The Great Pumpkin" is a fictional holiday figure in thecomic stripPeanuts byCharles M. Schulz.[142]
C. foetidissima contains asaponin that can be obtained from the fruit and root. This can be used as a soap, shampoo, and bleach. Prolonged contact can cause skin irritation.[143][144] Pumpkin is also used in cosmetics.[145]
Cucurbita have been used in various cultures as folk remedies. Pumpkins have been used byNative Americans to treat intestinal worms and urinary ailments. This Native American remedy was adopted by American doctors in the early nineteenth century as ananthelmintic for the expulsion of worms.[146] In southeastern Europe, seeds ofC. pepo were used to treat irritable bladder andbenign prostatic hyperplasia.[147] In Germany, pumpkin seed is approved for use by theCommission E, which assesses folk and herbal medicine, for irritated bladder conditions andmicturition problems of prostatic hyperplasia stages 1 and 2, although the monograph published in 1985 noted a lack of pharmacological studies that could substantiate empirically found clinical activity.[148] TheFDA in the United States, on the other hand, banned the sale of all such non-prescription drugs for the treatment of prostate enlargement in 1990.[149]
Halloween is widely celebrated with jack-o-lanterns made of large orange pumpkins carved with ghoulish faces and illuminated from inside with candles.[178] The pumpkins used for jack-o-lanterns areC. pepo,[179][180] not to be confused with the ones typically used for pumpkin pie in the United States, which areC. moschata.[122]Kew Gardens marked Halloween in 2013 with a display of pumpkins, including a towering pyramid made of many varieties of squash, in the Waterlily House during its "IncrEdibles" festival.[155]
^Due to wide variation in how the terms squash, pumpkin, and gourd are used, even among academics, in this article, the term squash can refer to any member of the genusCucurbita. Pumpkin and gourd are used to refer to species, varieties, and cultivars commonly referred to by those terms.[4]
^Ferriol, María; Picó, Belén (2007). "3".Handbook of Plant Breeding: Vegetables I. New York:Springer. p. 317.ISBN978-0-387-72291-7.The common terms "pumpkin", "squash", "gourd", "cushaw", "ayote", "zapallo", "calabaza", etc. are often applied indiscriminately to different cultivated species of the New World genusCucurbita L. (Cucurbitaceae):C. pepo L.,C. maxima Duchesne,C. moschata Duchesne,C. argyrosperma C. Huber andC. ficifolia Bouché.
^abcdDecker-Walters, Deena S.; Staub, Jack E.; Chung, Sang-Min; Nakata, Eijiro; Quemada, Hector D. (2002). "Diversity in Free-Living Populations ofCucurbita pepo (Cucurbitaceae) as Assessed by Random Amplified Polymorphic DNA".Systematic Botany.27 (1).American Society of Plant Taxonomists:19–28.JSTOR3093892.
^Millán, R. (1945). "Variaciones del Zapallito AmargoCucurbita andreana y el Origen deCucurbita maxima".Revista Argentina de Agronomía (in Spanish).12:86–93.
^Schrager, Victor (2004).The Compleat Squash: A Passionate Grower's Guide to Pumpkins, Squash, and Gourds. New York: Artisan. p. 25.ISBN978-1-57965-251-7.
^Hurd, Paul D.; Linsley, E. Gorton (1971). "Squash and Gourd Bees (Peponapis,Xenoglossa) and the Origin of the CultivatedCucurbita".Evolution.25 (1). St. Louis, MO: Society for the Study of Evolution:218–234.doi:10.2307/2406514.JSTOR2406514.PMID28562933.
^Whitaker, Thomas W.; Bemis, W. P. (1964). "Evolution in the GenusCucurbita".Evolution.18 (4):553–559.doi:10.2307/2406209.JSTOR2406209.
^abWinsor, J. A.; Davis, L. E.; Stephenson, A. G. (1987). "The Relationship Between Pollen Load and Fruit Maturation and the Effect of Pollen Load on Offspring Vigor inCucurbita pepo".The American Naturalist.129 (5):643–656.doi:10.1086/284664.JSTOR2461727.S2CID84901190.
^abRobinson, Richard W. (2000). "Rationale and Methods for Producing Hybrid Cucurbit Seed".Journal of New Seeds.1 (3–4):1–47.doi:10.1300/J153v01n03_01.S2CID85158524.
^Stephenson, Andrew G.; Devlin, B.; Horton, J. Brian (1988). "The Effects of Seed Number and Prior Fruit Dominance on the Pattern of Fruit Production inCucurbita pepo (Zucchini Squash)".Annals of Botany.62 (6):653–661.doi:10.1093/oxfordjournals.aob.a087705.
^Menezes, C. B.; Maluf, W. R.; Azevedo, S. M.; Faria, M. V.; Nascimento, I. R.; Gomez, L. A.; Bearzoti, E. (March 2005). "Inheritance of Parthenocarpy in Summer Squash (Cucurbita pepo L.)".Genetics and Molecular Research.4 (1):39–46.PMID15841434.
^Krupnick, Gary A.; Brown, Kathleen M.; Stephenson, Andrew G. (1999). "The Influence of Fruit on the Regulation of Internal Ethylene Concentrations and Sex Expression inCucurbita texana".International Journal of Plant Sciences.160 (2):321–330.doi:10.1086/314120.S2CID85794143.
^Murray, M. (1987). "Field Applications Of Ethephon For Hybrid And Open-Pollinated Squash (Cucurbita Spp) Seed Production".Acta Horticulturae.201 (201):149–156.doi:10.17660/ActaHortic.1987.201.15.
^"Plant Hormones". Charles Sturt University. Archived fromthe original on January 16, 2014. RetrievedJanuary 15, 2014.
^Wilson, Mack A.; Splittstoesser, Walter E. (1980). "The Relationship Between Embryo Axis Weight and Reserve Protein During Development and Pumpkin Seed Germination".Journal of Seed Technology.5 (2):35–41.JSTOR23432821.
^Oliver, Lawrence R.; Harrison, Steve A.; McClelland, Marilyn (1983). "Germination of Texas Gourd (Cucurbita texana) and Its Control in Soybeans (Glycine max)".Weed Science.31 (5):700–706.doi:10.1017/S0043174500070211.JSTOR4043694.S2CID182243467.
^Horak, Michael J.; Sweat, Jonathan K. (1994). "Germination, Emergence, and Seedling Establishment of Buffalo Gourd (Cucurbita foetidissima)".Weed Science.42 (3):358–363.doi:10.1017/S0043174500076621.JSTOR4045510.S2CID132074382.
^Beecroft, Penny; Lott, John N. A. (1996). "Changes in the Element Composition of Globoids FromCucurbita maxima andCucurbita andreana Cotyledons During Early Seedling Growth".Canadian Journal of Botany.74 (6):838–847.Bibcode:1996CaJB...74..838B.doi:10.1139/b96-104.
^Subin, M. P.; Francis, Steffy (2013). "Phytotoxic Effects of Cadmium on Seed Germination, Early Seedling Growth and Antioxidant Enzyme Activities inCucurbita maxima Duchesne".International Research Journal of Biological Sciences.2 (9):40–47.Bibcode:1996CaJB...74..838B.doi:10.1139/b96-104.
^Fenner, G. P.; Patteron, G. W.; Lusby, W. R. (1989). "Developmental Regulation of Sterol Biosynthesis inCucurbita maxima L.".Lipids.24 (4):271–277.doi:10.1007/BF02535162.S2CID37220982.
^abLinnaeus, Carl (1753)."Cucurbita".Species Plantarum. Vol. 2. Stockholm: Impensis Laurentii Salvii via Biodiversity Heritage Library. p. 1010.Archived from the original on July 2, 2017. RetrievedFebruary 20, 2018.
^Natural History Museum (2022)."Cucurbita".The Linnaean Plant Name Typification Project. Natural History Museum.doi:10.5519/qwv6u7j5.Archived from the original on November 29, 2014. RetrievedNovember 4, 2013.
^"Cucurbita".The Plant List.Archived from the original on March 17, 2019. RetrievedJanuary 1, 2015.
^abcWhitaker, T.W.; Bemis, W.P. (1975). "Origin and Evolution of the Cultivated Cucurbita".Bulletin of the Torrey Botanical Club.102 (6):362–368.doi:10.2307/2484762.JSTOR2484762.
^Janssen, Don (August 14, 2006)."Curbit Family & Cross-Pollination". University of Nebraska – Lincoln.Archived from the original on February 13, 2015. RetrievedJanuary 14, 2015.
^abBemis, W. P.;Whitaker, Thomas W. (April 1969). "The XerophyticCucurbita of Northwestern Mexico and Southwestern United States".Madroño.20 (2). California Botanical Society:33–41.JSTOR41423342.
^Kubitzki, Klaus (2011).Flowering Plants. Eudicots: Sapindales, Cucurbitales, Myrtaceae. Heidelberg: Springer Science & Business Media. pp. 120–122.ISBN978-3-642-14397-7.The fossil record of Cucurbitaceae and indeed of the order Cucurbitales is sparse.. The oldest fossils are seeds from the Uppermost Paleocene and Lower Eocene London Clay (65MA)..Bryonia-like seeds from fossil beda at Tambov, Western Siberia date to the Lower Sarmat, 15–13 MA ago. Subfossil records ofCucurbita pepo have been dated to 8,000–7,000 B.C. at Guila Naquitz ..., those ofC. moschata in the northern Peruvian Andes to up to 9,200 B.P.
^abcWilson, Hugh D."What isCucurbita texana?".Free-living Cucurbita pepo in the United States Viral Resistance, Gene Flow, and Risk Assessment. Texas A&M Bioinformatics Working Group. Archived fromthe original on September 27, 2013. RetrievedSeptember 8, 2013.
^abcdefghiHeistinger, Andrea (2013).The Manual of Seed Saving: Harvesting, Storing, and Sowing Techniques for Vegetables, Herbs, and Fruits. Portland, OR: Timber Press. p. 278.ISBN978-1-60469-382-9.
^abDecker, Deena S.; Wilson, Hugh D. (1987). "Allozyme Variation in theCucurbita pepo Complex:C. pepo var.ovifera vs.C. texana".Systematic Botany.12 (2). American Society of Plant Taxonomists:263–273.doi:10.2307/2419320.JSTOR2419320.
^"Squash". Texas A&M University.Archived from the original on October 29, 2013. RetrievedOctober 26, 2013.
^Salama, El-Sayed A.; Sill, W. H. Jr. (1968). "Resistance to Kansas Squash Mosaic Virus Strains Among Cucurbita Species".Transactions of the Kansas Academy of Science.71 (1):62–68.doi:10.2307/3627399.JSTOR3627399.
^"Cucurbita digitata A. Gray". United States Department of Agriculture.Archived from the original on September 21, 2013. RetrievedSeptember 1, 2013.
^"Cucurbita ficifolia Bouché". United States Department of Agriculture.Archived from the original on September 21, 2013. RetrievedSeptember 1, 2013.
^Mihranian, Valentine H.; Abou-Chaar, Charles I. (1968). "Extraction, Detection, and Estimation of Cucurbitin inCucurbita Seeds".Lloydia.31 (1). American Society of Pharmacognosy:23–29.
^Assessment report onCucurbita pepo L.(PDF) (Report). Committee on Herbal Medicinal Products (HMPC), European Medicines Agency. September 13, 2011. pp. 25–26. Archived fromthe original(PDF) on November 22, 2015. RetrievedNovember 21, 2015.
^Preedy, Victor R.; Watson, Ronald Ross; Patel, Vinwood B. (2011).Nuts and Seeds in Health and Disease Prevention. London: Academic Press. p. 936.ISBN978-0-12-375688-6.
^Barbieri, L.; Polito, L.; Bolognesi, A.; Ciani, M.; Pelosi, E.; Farini, V.; Jha, A. K.; Sharma, N.; Vivanco, J. M.; Chambery, A.; Parente, A.; Stirpe, F. (May 2006). "Ribosome-inactivating Proteins in Edible Plants and Purification and Characterization of a New Ribosome-inactivating Protein FromCucurbita moschata".Biochimica et Biophysica Acta (BBA) - General Subjects.1760 (5):783–792.doi:10.1016/j.bbagen.2006.01.002.PMID16564632.
^abChen, Jian Chao; Chiu, Ming Hua; Nie, Rui Lin; Cordell, Geoffrey A.; Qui, Samuel X. (2005). "Cucurbitacins and Cucurbitane Glycosides: Structures and Biological Activities".Natural Product Reports.22 (5):386–399.doi:10.1039/B418841C.PMID16010347.
^Mansour, Esam H.; Dworschák, Erno; Lugasi, Andrea; Barna, Barna; Gergely, Anna (1993). "Nutritive Value of Pumpkin (Cucurbita Pepo Kakai 35) Seed Products".Journal of the Science of Food and Agriculture.61 (1):73–78.Bibcode:1993JSFA...61...73M.doi:10.1002/jsfa.2740610112.
^Lazos, E. S. (July 1992). "Certain Functional Properties of Defatted Pumpkin Seed Flour".Plant Foods for Human Nutrition.42 (3):257–273.doi:10.1007/bf02193934.PMID1502127.S2CID1809751.
^Geisler, Malinda (May 2012)."Squash". Agricultural Marketing Resource Center, Iowa State University.Archived from the original on September 29, 2013. RetrievedOctober 13, 2013.
^abRichardson, R. W."Squash and Pumpkin"(PDF). United States Department of Agriculture, Agricultural Research Service, National Plant Germplasm System. Archived fromthe original(PDF) on September 24, 2015. RetrievedSeptember 2, 2013.
^BerrinLarco Museum, Katherine (1997).The Spirit of Ancient Peru: Treasures from the Museo Arqueológico Rafael Larco Herrera. New York: Thames and Hudson.ISBN978-0-500-01802-6.
^Cutler, H.C.; Whitaker, T.W. (1961). "History and Distribution of the Cultivated Cucurbits in the Americas".American Antiquity.26 (4):469–485.doi:10.2307/278735.JSTOR278735.S2CID161495351.(subscription required)
^Heiser, Charles B. (2016).The Gourd Book. Norman: University of Oklahoma Press. pp. 9,33–34.ISBN978-0806155678.
^Xiao, S. H.; Keiser, J.; Chen, M. G.; Tanner, M.; Utzinger, J. (2010). "Research and Development of Antischistosomal Drugs in the People's Republic of China a 60-year review".Advances in Parasitology.73:231–295.doi:10.1016/S0065-308X(10)73009-8.ISBN9780123815149.PMID20627145.
^Andrade-Cetto, A.; Heinrich, M. (July 2005). "Mexican Plants With Hypoglycaemic Effect Used in the Treatment of Diabetes".Journal of Ethnopharmacology.99 (3):325–348.doi:10.1016/j.jep.2005.04.019.PMID15964161.
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