Cucurbita
Squash | |
---|---|
Cucurbita fruits come in an assortment of colors and sizes. | |
Cross section of Cultivated Cucurbita of Canada | |
Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Clade: | Angiosperms |
Clade: | Eudicots |
Clade: | Rosids |
Order: | Cucurbitales |
Family: | Cucurbitaceae |
Tribe: | Cucurbiteae |
Genus: | Cucurbita L. |
Synonyms[1] | |
Cucurbita (
Most Cucurbita species are herbaceous vines that grow several meters in length and have tendrils, but non-vining "bush" cultivars of C. pepo and C. maxima have also been developed. The yellow or orange flowers on a Cucurbita plant are of two types: female and male. The female flowers produce the fruit and the male flowers produce pollen. Many North and Central American species are visited by specialist bee pollinators, but other insects with more general feeding habits, such as honey bees, also visit.
There is debate about the taxonomy of the genus and the number of accepted species varies from 13 to 30. The five domesticated species are Cucurbita argyrosperma, C. ficifolia, C. maxima, C. moschata, and C. pepo, all of which can be treated as winter squash because the full-grown fruits can be stored for months. However, C. pepo includes some cultivars that are better used only as summer squash.
The fruits of the genus Cucurbita are good sources of nutrients, such as vitamin A and vitamin C, among other nutrients according to species. The fruits have many culinary uses including pumpkin pie, biscuits, bread, desserts, puddings, beverages, and soups. Although botanical fruits, Cucurbita gourds such as squash are typically cooked and eaten as vegetables. Pumpkins see more varied use, and are eaten both as vegetables and as desserts such as pumpkin pie.
Description
Cucurbita species fall into two main groups. The first group consists of
The typical cultivated Cucurbita species has five-lobed or
The species are
Cucurbita fruits are large and fleshy.
Reproductive biology
All species of Cucurbita have 20 pairs of chromosomes.[16] Many North and Central American species are visited by specialist pollinators in the apid tribe Eucerini, especially the genera Peponapis and Xenoglossa, and these squash bees can be crucial to the flowers producing fruit after pollination.[6][17][18]
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,[19] an effect called xenia. Competitively grown specimens are therefore often hand-pollinated to maximize the number of seeds in the fruit, which increases the fruit size; this pollination requires skilled technique.[20][21] Seedlessness is known to occur in certain cultivars of C. pepo.[22][23]
The most 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.[24] The plant hormones ethylene and auxin are key in fruit set and development.[25] 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",[24] and male flowers are more frequent, an effect that appears due to reduced natural ethylene production within the plant stem.[26] 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.[20][27]
The plant hormone gibberellin, 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]
Germination and seedling growth
Seeds with maximum germination potential develop (in C. moschata) by 45 days after anthesis, and seed weight reaches its maximum 70 days after anthesis.[30] Some varieties of C. pepo germinate best with eight hours of sunlight daily and a planting depth of 1.2 centimeters (1⁄2 in). Seeds planted deeper than 12.5 centimeters (5 in) are not likely to germinate.[31] In C. 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.
Taxonomy
Cucurbita was formally described in a way that meets the requirements of modern
The Cucurbita digitata, C. foetidissima, C. galeotti, and C. pedatifolia
Various taxonomic treatments have been proposed for Cucurbita, ranging from 13 to 30 species.[3] 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:[6][42][43][44]
- synonymC. mixta) – cushaw pumpkin; origin: Mexico
- C. kellyana, origin: Pacific coast of western Mexico
- C. palmeri, origin: Pacific coast of northwestern Mexico
- C. sororia, origin: Pacific coast Mexico to Nicaragua, northeastern Mexico
- C. digitata – fingerleaf gourd; origin: southwestern United States (USA), northwestern Mexico
- C. ecuadorensis, origin: Ecuador's Pacific coast
- C. ficifolia – figleaf gourd, chilacayote, alcayota; origin: Mexico, Panama, northern Chile and Argentina
- C. foetidissima – stinking gourd, buffalo gourd; origin: Mexico
- C. galeottii, little known; origin: Oaxaca, Mexico
- C. lundelliana, origin: Mexico, Guatemala, Belize
- C. maxima – winter squash, pumpkin; origin: Argentina, Bolivia, Ecuador
- C. andreana, origin – Argentina
- C. moschata – butternut squash, 'Dickinson' pumpkin, golden cushaw; origin: Bolivia, Colombia, Ecuador, Mexico, Panama, Puerto Rico, Venezuela
- C. okeechobeensis, origin: Florida
- C. martinezii, origin: Mexican Gulf Coast and foothills
- C. pedatifolia, origin: Querétaro, Mexico
- C. moorei
- C. pepo – field pumpkin, summer squash, zucchini, vegetable marrow, courgette, acorn squash; origin: Mexico, USA
- C. fraterna, origin: Tamaulipas and Nuevo León, Mexico
- C. texana, origin: Texas, USA
- C. radicans – calabacilla, calabaza de coyote; origin: Central Mexico
- C. gracilior
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 several phenotypic 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:[16][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 as C. 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, and C. ecuadorensis were placed in their own separate species groups as they were not considered significantly close to any of the other species studied.
Phylogeny
The full phylogeny of this genus is unknown, and research was ongoing in 2014.[48][49] The following cladogram of Cucurbita phylogeny is based upon a 2002 study of mitochondrial DNA by Sanjur and colleagues.[50]
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Distribution and habitat
This section may require copy editing for redundancy. (December 2023) |
The ancestral species of the genus Cucurbita were present in the Americas before the
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.[6] Of the 27 species that Nee delineates, five are domesticated. Four of these, C. argyrosperma, C. ficifolia, C. moschata, and C. pepo, originated and were domesticated in Mesoamerica; the fifth, C. maxima, originated and was domesticated in South America.[9]
Within C. pepo, the pumpkins, the scallops, and possibly the crooknecks are ancient and were domesticated at different times and places. The domesticated forms of C. 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 of C. 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. argyrosperma is not as widespread as the other species. The wild form C. a. subsp. sororia is found from Mexico to Nicaragua, and cultivated forms are used in a somewhat wider area stretching from Panama to the southeastern United States.[9] It was probably bred for its seeds, which are large and high in oil and protein, but its flesh is of poorer quality than that of C. moschata and C. 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.[9]
C. ficifolia and C. moschata were originally thought to be Asiatic in origin, but this has been disproven. The origin of C. 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.[9]
C. maxima originated in South America over 4,000 years ago,[50] probably in Argentina and Uruguay. The plants are sensitive to frost, 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.[6] Types of C. maxima include triloba,[59] zapallito,[60] zipinka,[61] Banana, Delicious, Hubbard, Marrow (C. maxima Marrow), Show, and Turban.[62]
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 to C. 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).[9] Groups of C. 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 being
Cultivar group | Botanical name | Image | Description |
---|---|---|---|
Acorn | C. pepo var. turbinata | 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[9][70][71] | |
Cocozzelle | C. pepo var. Ionga | Summer squash, long round slender fruit that is slightly bulbous at the apex,[69] similar to fastigata, ex: Cocozelle von tripolis[9][70][71] | |
Crookneck | C. pepo var. torticollia (also torticollis) | 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[9][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;[9][70][71] includes C. pepo subsp. pepo var. styriaca, used for Styrian pumpkin seed oil[73] | |
Scallop | C. pepo var. clypeata; called C. melopepo by Linnaeus[7] | Summer squash, prefers half-shrubby habitat, flattened or slightly discoidal shape, with undulations or equatorial edges,[69] ex: Pattypan squash[9][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[9][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)[9][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 to fastigata, ex: Zucchini[9][70][71] | |
Ornamental gourds | C. pepo var. ovifera | Non-edible,[72] field squash closely related to C. texana, vine habitat, thin stems, small leaves, three sub-groups: C. pepo var. ovifera (egg-shaped, pear-shaped), C. pepo var. aurantia (orange color), and C. 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] |
Ecology
Cucurbita species are used as food plants by the larvae of some Lepidoptera species, including the cabbage moth (Mamestra brassicae), Hypercompe indecisa, and the turnip moth (Agrotis segetum).[74] Cucurbita can be susceptible to the pest Bemisia argentifolii (silverleaf whitefly)[75] as well as aphids (Aphididae), cucumber beetles (Acalymma vittatum and Diabrotica undecimpunctata howardi), squash bug (Anasa tristis), the squash vine borer (Melittia cucurbitae), and the two-spotted spidermite (Tetranychus urticae).[76] The squash bug causes major damage to plants because of its very toxic saliva.[77] The
Species in the genus Cucurbita are susceptible to some types of mosaic virus including: cucumber mosaic virus (CMV), papaya ringspot virus-cucurbit strain (PRSV), squash mosaic virus (SqMV), tobacco ringspot virus (TRSV),[80] watermelon mosaic virus (WMV), and zucchini yellow mosaic virus (ZYMV).[81][82][83][84] PRSV is the only one of these viruses that does not affect all cucurbits.[81][85] SqMV and CMV are the most common viruses among cucurbits.[86][87] Symptoms of these viruses show a high degree of similarity, which often results in laboratory investigation being needed to differentiate which one is affecting plants.[80]
Cultivation
History
The genus was part of the culture of almost every native peoples group from southern South America to southern Canada.
The earliest known evidence of the domestication of Cucurbita dates back at least 8,000 years ago, predating the domestication of other crops such as
Squash was domesticated first, followed by maize and then beans, becoming part of the
Production
Squash and gourd* production (2021) | |
---|---|
Country | Production (millions of tonnes) |
China | 7.4 |
Ukraine | 1.3 |
Russia | 1.2 |
United States | 1.1 |
Turkey | 0.8 |
Mexico | 0.7 |
Italy | 0.6 |
Indonesia | 0.5 |
Egypt | 0.4 |
World | 23.4 |
*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.
Toxicity
Cucurbitacin is a plant steroid present in wild Cucurbita and in each member of the family Cucurbitaceae. Poisonous to mammals,[79] it is found in quantities sufficient to discourage herbivores. It makes wild Cucurbita and most ornamental gourds, with the exception of an occasional C. fraterna and C. sororia, bitter to taste.[3][67][108] Ingesting too much cucurbitacin can cause stomach cramps, diarrhea and even collapse.[109] This bitterness is especially prevalent in wild Cucurbita; 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,[3] there are occasional reports of cucurbitacin causing illness in humans.[3] Cucurbitacin is also used as a lure in insect traps.[108]
Uses
Nutrition
Nutritional value per 100 g (3.5 oz) | |
---|---|
Energy | 69 kJ (16 kcal) |
3.4 g | |
Sugars | 2.2 g |
Dietary fiber | 1.1 g |
0.2 g | |
1.2 g | |
Niacin (B3) | 3% 0.487 mg |
Pantothenic acid (B5) | 3% 0.155 mg |
Vitamin B6 | 13% 0.218 mg |
Folate (B9) | 7% 29 μg |
Vitamin C | 19% 17 mg |
Vitamin K | 3% 3 μg |
Minerals | Quantity %DV† |
Iron | 2% 0.35 mg |
Magnesium | 4% 17 mg |
Manganese | 8% 0.175 mg |
Phosphorus | 3% 38 mg |
Potassium | 6% 262 mg |
Zinc | 3% 0.29 mg |
Other constituents | Quantity |
Water | 95 g |
Link to USDA Database entry, for comparison, see values for raw pumpkin | |
†Percentages estimated using US recommendations for adults.[111] |
As an example of Cucurbita, raw summer squash is 94% water, 3%
Culinary
The family Cucurbitaceae has many species used as human food.
Squashes are primarily grown for the fresh food market.[120]
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 the
In India, squashes (ghiya) are cooked with seafood such as prawns.
In culture
Art, music, and literature
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.
Though native to the western hemisphere, Cucurbita began to spread to other parts of the world after
In 1952, Stanley Smith Master, using the pen name Edrich Siebert, wrote "The Marrow Song (Oh what a beauty!)" to a tune in
8 time. It became a popular hit in Australia in 1973,[137] and was revived by the Wurzels in Britain on their 2003 album Cutler of the West.[138][139] John Greenleaf Whittier wrote a poem entitled The Pumpkin in 1850.[140] "The Great Pumpkin" is a fictional holiday figure in the comic strip Peanuts by Charles M. Schulz.[141]
Cleansing and personal care uses
C. foetidissima contains a saponin 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.[142][143] Pumpkin is also used in cosmetics.[144]
Folk remedies
Cucurbita have been used in various cultures as folk remedies. Pumpkins have been used by
In China,
In Mexico, herbalists use C. ficifolia in the belief that it reduces
Festivals
Cucurbita fruits including pumpkins and marrows are celebrated in festivals in countries such as Argentina, Austria,
Halloween is widely celebrated with jack-o-lanterns made of large orange pumpkins carved with ghoulish faces and illuminated from inside with candles.[176] The pumpkins used for jack-o-lanterns are C. pepo,[177][178] not to be confused with the ones typically used for pumpkin pie in the United States, which are C. moschata.[121] 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.[179]
See also
- List of gourds and squashes in the genus Cucurbita
- List of squash and pumpkin dishes
Notes
References
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The common terms "pumpkin", "squash", "gourd", "cushaw", "ayote", "zapallo", "calabaza", etc. are often applied indiscriminately to different cultivated species of the New World genus Cucurbita L. (Cucurbitaceae): C. pepo L., C. maxima Duchesne, C. moschata Duchesne, C. argyrosperma C. Huber and C. ficifolia Bouché.
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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 of Cucurbita pepo have been dated to 8,000-7,000 B.C. at Guila Naquitz ..., those of C. moschata in the northern Peruvian Andes to up to 9,200 B.P.
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