Cucurbita

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Cucurbita
Squashes at Kew Gardens IncrEdibles 2013.jpg
Cucurbita fruits come in an assortment of colors and sizes
Scientific classification
Kingdom: Plantae
(unranked): Angiosperms
(unranked): Eudicots
(unranked): Rosids
Order: Cucurbitales
Family: Cucurbitaceae
Subfamily: Cucurbitoideae
Tribe: Cucurbiteae
Genus: Cucurbita
L.
Synonyms[2]

Cucurbita (Latin for gourd)[3] is a genus in the gourd family Cucurbitaceae native to and first cultivated in the Andes and Mesoamerica. The Cucurbita genus is an important source of human food, beverages, medicine, oil, and traditionally of detergent. Some Cucurbita species were brought to Europe after the discovery of America and are now used in many parts of the world. The plants, referred to as squash, pumpkin or gourd depending on species, variety, and local parlance,[a] are grown for their edible fruits and seeds. Gourds, also called bottle-gourds, which are used as utensils or vessels, are native to Africa and belong to the genus Lagenaria, which is in the same family and subfamily as Cucurbita but in a different tribe.

There are five domesticated species: Cucurbita argyrosperma, C. ficifolia, C. maxima, C. moschata, and C. pepo. All of these species can be treated as winter squash because the full-grown fruits can be stored for months; C. pepo includes some cultivars that are better used only as summer squash. There is no universal agreement on how to handle the taxonomy of the genus, with the number of species listed varying from 13 to 30. There is uncertainty as to the ancestry of some of the species that have been cultivated for millennia.

Most Cucurbita species are vines that grow several meters in length and have tendrils, but non-vining "bush" cultivars of C. pepo and C. maxima have been developed. The flowers are yellow or orange, and there are two types of flowers on a Cucurbita plant: the female flowers that produce the fruit and the male flowers that 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. The fruits of the Cucurbita genus are good sources of several nutrients such as vitamin A, vitamin C, dietary fiber, niacin, folic acid, and iron, and like all plant products are free of cholesterol. The plants contain the toxins cucurbitin, cucurmosin, and cucurbitacin.

Cucurbita fruits have played a role in human culture for at least 2,000 years. They are often represented in Moche ceramics from Peru. After Christopher Columbus's arrival in the New World, paintings of squashes started to appear in Europe early in the sixteenth century. Among other uses, Cucurbita seeds have been employed in traditional medicine, and extracts are used in cosmetics for dry and sensitive skin. The fruits have many culinary uses including pumpkin pie, biscuits, bread, desserts, puddings, and soups. Pumpkins and other Cucurbita fruits are celebrated in festivals such as Halloween, pumpkin chucking, the Keene Pumpkin Fest, and flower and vegetable shows in many countries.

Description[edit]

The leaves of Cucurbita moschata often have white spots near the veins

Most Cucurbita species are climbing annual vines and are mesophytes, plants which require a more or less continuous water supply. The perennial species grow in tropical zones and are xerophytes, plants which tolerate dry conditions well. Growing 5 to 15 meters (16 to 49 ft), the plant stem produces tendrils to help it climb adjacent plants and structures or along the ground. Most species do not readily root from the nodes; a notable exception is Cucurbita ficifolia, and the four other cultivated mesophytes do this to a lesser extent. The vine of the perennial Cucurbita can become semiwoody if left to grow. There is wide variation in size, shape, and color among Cucurbita fruits, and even within a single species. C. ficifolia is an exception to this, being highly uniform in appearance.[5] The morphological variation in the species C. pepo[6] and C.  maxima[7] are so vast that its various subspecies and cultivars have been misidentified as totally separate species.

The typical cultivated Cucurbita species has five-lobed or palmately divided leaves with long petioles, and the leaves alternately arranged on the stem. The stems in some but not all species are angular. All of the above-ground parts may be hairy with various types of trichomes, which are often hardened and sharp. Spring-like tendrils grow from each node and are branching in some species. C. argyrosperma has ovate-cordate (egg-shaped to heart-shaped) leaves. The shape of C. pepo leaves varies widely. C. moschata plants can have light or dense pubescence. 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]

There are male (staminate) and female (pistillate) flowers (unisexual flowers) on a single plant (monoecious), and these grow singly, appearing from the leaf axils. Flowers have five fused yellow to orange petals (the corolla) and a green bell-shaped calyx. Male flowers in Cucurbitaceae generally have five stamens, but in Cucurbita there are only three, and their anthers are joined together so that there appears to be one.[9][10] Female flowers have thick pedicels, and an inferior ovary with 3–5 stigmas that each have two lobes.[11][8] The female flowers of C. argyrosperma and C. ficifolia have larger corollas than the male flowers.[8] Female flowers of C. pepo have a small calyx, but the calyx of C. moschata male flowers is comparatively short.

Cucurbita fruits are large and fleshy.[9] Botanists classify the Cucurbita fruit as a pepo, which is a special type of berry derived from an inferior ovary, with a thick outer wall or rind with hypanthium tissue forming an exocarp around the ovary, and a fleshy interior composed of mesocarp and endocarp. The term "pepo" is used primarily for Cucurbitaceae fruits, where this fruit type is common, but the fruits of Passiflora and Carica are sometimes also considered pepos.[12][13][14] Seeds are attached to the ovary wall (parietal placentation) and not to the center. Seeds are large and fairly flat with an embryo which has two rather large cotyledons.[11] Wild fruit specimens can be as small as 4 centimeters (1.6 in), but certain domesticated specimens can weigh well over 300 kilograms (660 lb).[8] The current world record was set in October 2014 by John Hawkley of Napa County, California with a 933.5-kilogram (2,058 lb) pumpkin.[15]

Reproductive biology[edit]

Cucurbita female flower with pollinating squash bees

Every species of Cucurbita has 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.[5][17] Competitively grown specimens are often hand-pollinated to maximize the number of seeds in the fruit, which increases the fruit size (an effect called xenia); this pollination requires proper technique.[18] Seedlessness is known to occur in certain cultivars of C. pepo.[19][20]

Male flower, part of the perianth and 1 filament removed

The most critical factors in flowering and fruit set are physiological, not climatic, factors.[21] The plant hormones ethylene and auxin are key in fruit set and development.[22] 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",[21] and male flowers are more frequent, an effect that appears to be due to reduced natural ethylene production within the plant stem.[23] 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.[24]

Gibberellin produced in the stamens is a plant hormone essential for the development of all parts of the male flowers, but the development of female flowers is not yet understood.[25] Gibberellin is also involved in other developmental processes of plants such as seed and stem growth.[26]

Germination and seedling growth[edit]

Kabocha seedling 7 days after being sown

Seed germination in some species of Cucurbita has been directly linked to embryo axis weight and reserve protein. Seed with maximum germination potential in C. moschata had developed by 45 days after anthesis, and seed weight reached its maximum 70 days after anthesis.[27] Some varieties of C. pepo germinate best with eight hours of sunlight daily and a planting depth of 1.2 centimeters (0.47 in). Seeds planted deeper than 12.5 centimeters (4.9 in) are not likely to germinate.[28] 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% germination rate, over 90% of the plants had sprouted after 29 days from planting.[29]

High levels of pollen load are known to produce more seeds in fruits, larger fruits, greater likelihood of fruit maturation, and faster and more likely seed germination with larger seedlings.[30] Various combinations of light and mineral nutrients have a significant effect during the various stages of plant growth. These effects vary significantly between the different species of Cucurbita. Phytate forms in seeds tissues as spherical crystalline intrusions in protein bodies called globoids. The nutrients in globoids are eventually used completely during seedling growth.[31] Heavy metal contamination, including cadmium, has a significant negative impact on plant growth.[32] Cucurbita plants grown in the spring tend to grow larger than those grown in the autumn.[33]

Species[edit]

An assortment of fruits of C. maxima and C. pepo

Cucurbita was formally described in 1753 in Linnaeus's Species Plantarum, which established modern botanical nomenclature. Cucurbita pepo is the type species of the genus.[34][35] The Cucurbita digitata, C. foetidissima, C. galeotti, and C. pedatifolia species groups are xerophytes, 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][36] The five domesticated species are mostly isolated from each other by sterility barriers and have different physiological characteristics.[36] Some cross pollinations can occur: C. pepo with C. argyrosperma and C. moschata; and C. maxima with C. moschata. Cross pollination does occur readily within the family Cucurbitaceae.[37] The buffalo gourd (C. foetidissima), which does not taste good, has been used as an intermediary as it can be crossed with all the common Cucurbita.[11]

Various taxonomic treatments have been proposed for Cucurbita, ranging from 13–30 species.[3] In 1990, Michael Nee classified them into the following 13 species groups (27 species total), listed by group and alphabetically, with geographic origin:[5][38][39]

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][42]

  • 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 four separate species groups as they were not considered significantly close to any of the other species studied.

Phylogeny[edit]

The full phylogeny of this genus is unknown, and research is ongoing in 2014.[43][44] The following cladogram of Cucurbita phylogeny is based upon a 2002 study of mitochondrial DNA by Sanjur et al.[45]



Sechium edule




C. ficifolia




C. foetidissima





C. maxima and C. andreana



C. ecuadorensis





C. martinezii




C. pepo subspp. fraterna and ovifera




C. pepo subsp. pepo




C. sororia, in part




C. sororia, in part and C. argyrosperma











Habitat and distribution[edit]

A festival-winning 742 kilograms (1,636 lb) pumpkin in 2009

There are five domesticated species. Four of them, Cucurbita argyrosperma, C. ficifolia, C. moschata, and C. pepo, originated and were domesticated in Mesoamerica. For one of them, C. maxima, these events occurred in South America.[8]

Within C. pepo, the pumpkins, scallops, and possibly crooknecks are ancient and were domesticated separately. The domesticated forms of C. pepo have larger fruits and larger yet fewer seeds.[46] In a 1989 study on the origins and development of C. pepo, botanist Harry Paris suggested that the original wild specimen was 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 is a cross between the scallop and pumpkin.[46]

Not as widespread as the other species, Cucurbita argyrosperma is found from Mexico to Nicaragua. It has large seeds that are high in oil and protein and was probably bred for its seeds. Its flesh is of poorer quality than that of C. moschata and C. pepo. It is found from sea level to as high as 1,800 meters (5,900 ft) in dry areas or areas with a defined rainy season. Seeds are sown in May and June at the start of the rainy season.[8]

Cucurbita 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 altitudes from 1,000 meters (3,300 ft) to 3,000 meters (9,800 ft) in areas with heavy rainfall. It does not hybridize well with the other cultivated species as it has significantly different enzymes and chromosomes.

Cucurbita maxima originated in South America over 4,000 years ago,[45] probably in Argentina and Uruguay. The plants are sensitive to frost, prefer bright sunlight, and soil with a pH of 6.0 to 7.0.[47] Cucurbita 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 of C. maxima include Triloba,[48] Zapallito,[49] Zipinka,[50] Banana, Delicious, Hubbard, Marrow (C. maxima Marrow), Show, and Turban.[51]

C. moschata 'Butternut'
Fruit of the 'Tromboncino' cultivar of the Crookneck (C. moschata) Group are eaten either when very young, or as mature winter squash.

Cucurbita moschata is native to Latin America but the precise location of origin is uncertain.[52] 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 (7.9 to 11.8 in) wide. It generally grows at low altitudes in hot climates with heavy rainfall, but some varieties have been found above 2,200 meters (7,200 ft).[8] Groups of C. moschata include Cheese, Crookneck (C. moschata), and Bell.[51]

Cucurbita pepo is one of the oldest, if not the oldest, domesticated species with the oldest known locations being Oaxaca, Mexico, 8,000–10,000 years ago, and Ocampo, Tamaulipas, Mexico, about 7,000 years ago. It is known to have appeared in Missouri, United States, at least 4,000 years ago.[5][8][53][54] Debates about the origin of C. pepo have been on-going since at least 1857.[55] There have traditionally been two opposing theories about its origin: 1) that it is a direct descendant of C. texana and 2) that C. texana is merely feral C. pepo.[5] A more recent theory by botanist Thomas Andres in 1987 is that descendants of C. fraterna hybridized with C. texana,[56] resulting in two distinct domestication events in two different areas: one in Mexico and one in the eastern United States, with C. fraterna and C. texana, respectively, as the ancestral species.[8][45][56][57] C. pepo may have appeared in the Old World prior to 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 cm wide. All the subspecies, varieties, and cultivars are conspecific and interfertile.[6] In 1986 Paris proposed a revised taxonomy of the edible cultivated C. pepo with eight groups based on their basic shape and color, which varies widely.[46][58] All but a few C. pepo can be included in these groups.[58][8][59][60] There is one non-edible cultivated variety: C. pepo var. ovifera.[61]

A classification of cultivated C. pepo varieties based on Paris' eight groups and the one non-edible variety
Cultivar group Botanical name Image Description
Acorn C. pepo var. turbinata Starr 070730-7820 Cucurbita pepo.jpg 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,[58] ex: Acorn squash[8][59][60]
Cocozzelle C. pepo var. Ionga Cucurbita pepo Cocozelle fruits.jpg Summer squash, long round slender fruit that is slightly bulbous at the apex,[58] similar to fastigata, ex: Cocozelle von tripolis[8][59][60]
Crookneck C. pepo var. torticollia (also torticollis) Crooked Neck Squash.jpg 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,[58] ex: Yellow crookneck squash[8][59][60]
Pumpkin C. pepo var. pepo
Pumpkin 2 - Evan Swigart.jpg
Winter squash, creeping plant, round, oblate, or oval shape and round or flat on the ends,[58] ex: Pumpkin;[8][59][60] includes C. pepo subsp. pepo var. styriaca, used for Styrian pumpkin seed oil[62]
Scallop C. pepo var. clypeata; called C. melopepo by Linnaeus[6] Pattypan squash J1.jpg Summer squash, prefers half-shrubby habitat, flattened or slightly discoidal shape, with undulations or equatorial edges,[58] ex: Pattypan squash[8][59][60]
Straightneck C. pepo var. recticollis Cucurbita pepo Yellow Squash 3.jpg Summer squash, shrubby plant, with yellow or golden fruit and verrucose rind, similar to var. torticollia but a stem end that narrows,[58] ex: Yellow summer squash[8][59][60]
Vegetable marrow C. pepo var. fastigata Spaghetti Squash 700.jpg Summer and winter squashes, creeper traits and a semi-shrub, cream to dark green color, short round fruit with a slightly broad apex,[58] ex: Spaghetti squash (a winter variety)[8][59][60]
Zucchini C. pepo var. cylindrica Zucchini-Whole.jpg Summer squash, presently the most common group of cultivars, origin is recent (19th century), semi-shrubby, cylindrical fruit with a mostly consistent diameter,[58] similar to fastigata, ex: Zucchini[8][59][60]
Ornamental gourds C. pepo var. ovifera Cucurbita pepo var ovifera (crop).jpg Non-edible,[61] 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.[53]

Nutrients[edit]

Summer squash, all varieties, raw
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
Vitamins
Vitamin A equiv.
(1%)
10 μg
(1%)
120 μg
2125 μg
Thiamine (B1)
(4%)
0.048 mg
Riboflavin (B2)
(12%)
0.142 mg
Niacin (B3)
(3%)
0.487 mg
(3%)
0.155 mg
Vitamin B6
(17%)
0.218 mg
Folate (B9)
(7%)
29 μg
Vitamin C
(20%)
17 mg
Vitamin K
(3%)
3 μg
Trace metals
Iron
(3%)
0.35 mg
Magnesium
(5%)
17 mg
Manganese
(8%)
0.175 mg
Phosphorus
(5%)
38 mg
Potassium
(6%)
262 mg
Zinc
(3%)
0.29 mg
Other constituents
Water 95 g

Percentages are roughly approximated using US recommendations for adults.
Source: USDA Nutrient Database

Cucurbita have phytochemical constituents such as alkaloids, flavonoids, and palmitic, plus oleic and linoleic acids. Animal and in vitro experiments suggest that pumpkins have anti-diabetic, antioxidant, anticarcinogenic, and anti-inflammatory pharmacological properties.[63] Pumpkins and pumpkin seeds have high levels of crude protein, calcium, iron, potassium, phosphorus, magnesium, zinc,[64][65] and beta-carotene.[66] Cucurbita are good sources of vitamin A, vitamin C, potassium, dietary fiber, niacin, folic acid, and iron. They are free of cholesterol.

Toxins[edit]

Cucurbitin is an amino acid and a carboxypyrrolidine that is found in raw Cucurbita seeds.[67][68] Cucurbitin is an inhibitor of histidine decarboxylase, which is associated with inhibition of the biosynthesis of histamine, which in turn is responsible for the formation of the inflammatory response.[69] Cucurbitin causes degenerative changes in the reproductive organs of parasitic flukes.[70][71]

Cucurmosin is a ribosome inactivating protein found in the flesh and seed of Cucurbita,[72][73] notably Cucurbita moschata. Cucurmosin is more toxic to cancer cells than healthy cells.[72][74]

Cucurbitacin is a plant steroid poisonous to mammals found in wild Cucurbita in quantities sufficient to discourage herbivores,[75] and in each member of the Cucurbitaceae family.[76] It has a bitter taste, and is what makes wild Cucurbita and most ornamental gourds, with the exception of an occasional C. fraterna and C. sororia, bitter to taste.[3][56] Ingesting too much cucurbitacin can cause stomach cramps, diarrhea and even collapse.[77] Because of this bitterness that 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.[78] While the process of domestication has largely removed the bitterness from cultivated varieties,[3] there are occasional reports of cucurbitacin getting into the human food supply and causing illness.[3] Cucurbitacin is also used as a lure in insect traps.[76]

Pests and diseases[edit]

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).[79] Cucurbita can be susceptible to the pest Bemisia argentifolii (silverleaf whitefly)[80] 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 twospotted spidermite (Tetranychus urticae).[81] The squash bug causes major damage to plants because of its very toxic saliva.[82] Cucurbits, which are all members of the family Cucurbitaceae, are susceptible to diseases such as bacterial wilt (Erwinia tracheiphila), anthracnose (Colletotrichum spp.), fusarium wilt (Fusarium spp.), phytophthora blight (Phytophthora spp. water molds), and powdery mildew (Erysiphe spp.).[81] Defensive responses to viral, fungal, and bacterial leaf pathogens do not involve cucurbitacin.[75]

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),[83] Watermelon mosaic virus (WMV), and Zucchini yellow mosaic virus (ZYMV).[84][85][86][87] PRSV is the only one of these viruses that does not affect all cucurbits.[84][88] SqMV and CMV are the most common viruses among cucurbits.[89][90] 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.[83]

Production[edit]

Squashes are primarily grown for the fresh food market.[91] The Food and Agriculture Organization of the United Nations (FAO) reported that the ranking of the top five squash-producing countries was stable from 2005–2009. Those countries are: China, India, Russia, the United States, and Egypt. By 2012, Iran had moved into the 5th slot, with Egypt falling to 6th. The top 10 countries in terms of metric tons of squashes produced are:[92]

Top ten squash producers — 2012[92]
Country Production
(metric tons)
China China 6,140,840
India India 4,424,200
Russia Russia 988,180
United States USA 778,630
Iran Iran 695,600
Egypt Egypt 658,234
Mexico Mexico 522,388
Ukraine Ukraine 516,900
Italy Italy 508,075
Turkey Turkey 430,402
Top 10 total 15,663,449

The only other countries that rank in the top 20 where squashes are native are Cuba, which ranks 14th with 347,082 metric tons, and Argentina, which ranks 17th, with 326,900 metric tons.[92] In addition to being the 4th largest producer of squashes in the world, the United States is the world's largest importer of squashes, importing 271,614 metric tons in 2011, 95% of that from Mexico. Within the United States, the states producing the largest amounts are Florida, New York, California, and North Carolina.[91]

This is how Cucurbita compares to several other major Cucurbitaceae crops in terms of crop tonnage harvested:

Top ten cucumber producers — 2012[92]
Country Production
(metric tons)
China China 40,710,200
Iran Iran 1,811,630
Turkey Turkey 1,739,190
Russia Russia 1,161,870
United States USA 883,360
Ukraine Ukraine 860,100
Spain Spain 682,900
Egypt Egypt 631,408
Japan Japan 587,800
Indonesia Indonesia 547,141
Top 10 total 49,075,599
Top ten watermelon producers — 2012[92]
Country Production
(metric tons)
China China 56,649,725
Turkey Turkey 3,683,100
Iran Iran 3,466,880
Brazil Brazil 1,870,400
United States USA 1,866,660
Egypt Egypt 1,637,090
Uzbekistan Uzbekistan 1,182,400
Russia Russia 1,151,580
Mexico Mexico 1,036,800
Algeria Algeria 946,200
Top 10 total 73,490,835

History and domestication[edit]

Cucurbita pepo subsp. texana, from Les Grandes Heures d'Anne de Bretagne, 1503–1508, f. 161, earliest depiction of cucurbits in Europe

The ancestral species of the genus Cucurbita were present in the Americas before the arrival of humans,[93][94] and are native to the New World. The likely center of origin is southern Mexico, spreading south into what is now known as Mesoamerica and north to what is now the southwestern United States.[93] Evolutionarily speaking, the genus is relatively recent in origin, and no species within the genus is entirely genetically isolated. Cucurbita moschata can intercross with all the others, though the hybrid offspring may not themselves be fertile unless they become polyploid.[36] The genus was part of the culture of almost every native peoples group from southern South America to southern Canada.[94] Modern-day cultivated Cucurbita are not found in the wild.[5] Genetic studies of the mitochondrial gene nad1 show there were at least six independent domestication events of Cucurbita separating domestic species from their wild ancestors.[45] Species native to North America include C. digitata (calabazilla),[95] and C. foetidissima (buffalo gourd),[96] C. palmata (coyote melon), and C. pepo.[5] Some species, such as C. digitata and C. ficifolia, are referred to as gourds. Gourds, also called bottle-gourds, which are used as utensils or vessels, belong to the genus Lagenaria and are native to Africa. Lagenaria are in the same family and subfamily as Cucurbita but in a different tribe.[97]

The earliest known evidence of the domestication of Cucurbita dates back 8,000–10,000 years ago, predating the domestication of other crops such as maize and beans in the region by about 4,000 years.[5][53][54][98] This evidence was found in the Guilá Naquitz cave in Oaxaca, Mexico, during a series of excavations in the 1960s and 1970s, possibly beginning in 1959.[99][100] Later, more accurate, dating using accelerator mass spectrometers provided more specific dates. Solid evidence of domesticated Cucurbita pepo was found in the Guilá Naquitz cave in the form of increasing rind thickness and larger peduncles in the newer stratification layers of the cave. By circa 8,000 years BP the C. pepo peduncles found are consistently more than 10 millimeters (0.39 in) thick. Wild Cucurbita peduncles are always below this 10 mm barrier. Changes in fruit shape and color indicate that intentional breeding of C. pepo had occurred by no later than 8,000 years BP.[11][101][102] During the same time frame, average rind thickness increased from 0.84 millimeters (0.033 in) to 1.15 millimeters (0.045 in).[103]

The process of domesticating Cucurbita took place over 5,000–6,500 years in Mesoamerica. Squash was domesticated first, followed by maize and then beans, becoming part of the Three Sisters agricultural system of companion planting.[104][105] The English word "squash" derives from askutasquash (a green thing eaten raw), a word from the Narragansett language, which was documented by Roger Williams, the founder of Rhode Island, in his 1643 publication A Key Into the Language of America.[106] Similar words for squash exist in related languages of the Algonquian family.[107][46]

In human culture[edit]

Art, music, and literature[edit]

Moche Squash Ceramic. 300 A.D. Larco Museum

Squash, along with maize and beans, has been depicted in the art work of the native peoples of the Americas for at least 2,000 years.[108][109] For example, cucurbita are often represented in Moche ceramics.[108][110]

Though native to the western hemisphere, Cucurbita began to spread to other parts of the world after Christopher Columbus's arrival in the New World in 1492.[111][112] Until recently, the earliest known depictions of this genus in Europe was of Cucurbita pepo in De Historia Stirpium Commentarii Insignes in 1542 by the German botanist Leonhart Fuchs, but in 1992, two paintings, one of C. pepo and one of C. maxima, painted between 1515 and 1518, were identified in festoons at Villa Farnesina in Rome.[113] Also, in 2001 depictions of this genus were identified in Grandes Heures of Anne of Brittany (Les Grandes Heures d'Anne de Bretagne), a French devotional book, an illuminated manuscript, created between 1503 and 1508. This book contains an illustration known as Quegourdes de turquie, which was identified by cucurbit specialists as C. pepo subsp. texana in 2006.[114]

In 1952, Stanley Smith Master, using the pen name Edrich Siebert, wrote "The Marrow Song (Oh what a beauty!)" to a tune in 6/8 time. It became a popular hit in Australia in 1973,[115] and was revived by the Wurzels in Britain on their 2003 album Cutler of the West.[116][117] John Greenleaf Whittier wrote a poem entitled The Pumpkin in 1850.[118] "The Great Pumpkin" is a fictional holiday figure in the comic strip Peanuts by Charles M. Schulz.[119]

Medicinal uses[edit]

Cucurbita seeds were used in traditional Chinese medicine to treat the parasitic diseases ascariasis[120] and schistosomiasis.[68][120][121] Cucurbita extracts are used in cosmetics for dry and sensitive skin.[122][123][124] C. ficifolia is used to make soft and mildly alcoholic drinks,[8] and high doses of C. ficifolia have been shown to be successful in reducing blood sugar levels.[3] The fruit pulp of some species, such as C. foetidissima, can be used as a soap or detergent,[3][125][126] and buffalo gourd oil, made from Cucurbita foetidissima, was also made into soap.[125] Cucurbita fruits are an important source for humans of carotenoids, vitamin A, and rhodopsin, all of which are important to eye health.[11][127]

Culinary uses[edit]

Pumpkin custard made from kabocha, a cultivated variant of C. maxima

The Cucurbitaceae family has many species used as human food.[8] Cucurbita is one of the most important of those, with the various species being prepared and eaten in many ways. Although the stems and skins tend to be more bitter than the rest of the flesh,[77][128] the fruits and seeds of cultivated varieties are quite edible and need little or no preparation. In addition, the flowers and young leaves and shoot tips can be consumed.[129] The seeds and fruits of most varieties can be stored for long periods of time.[5] In particular, the sweet-tasting winter varieties store well due to their thick, inedible skins. Summer squash have a thin, edible skin. The seeds of both types can be ground into a flour or meal,[130] roasted, eaten raw, made into pumpkin seed oil,[62] or otherwise prepared. Long before European contact, Cucurbita had been a major food source for the native peoples of the Americas, and the species became an important food for European settlers, including the Pilgrims, even featuring at the first Thanksgiving.[11] Commercially made pumpkin pie mix is most often made from varieties of C. moschata; 'Libby's Select' uses the Select Dickinson Pumpkin variety of C. moschata for its canned pumpkin.[131] 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,[132] salads, soups, and stuffing.[133] The xerophytic species are proving useful in the search for nutritious foods that grow well in arid regions.[134]

Festivals[edit]

Towering pyramid of squashes in the Waterlily House, Kew Gardens, 2013

Cucurbita fruits including pumpkins and marrows are celebrated in festivals in countries such as Argentina, Bolivia,[135] Britain, Canada,[136] France,[137][138][139] Germany, Italy,[140][141][142][143][144] Japan,[145] Peru,[146] Portugal, Spain,[147] Switzerland,[148] and the United States. Argentina holds an annual nation-wide pumpkin festival Fiesta Nacional del Zapallo ("Squashes and Pumpkins National Fest"), in Ceres, Santa Fe,[149] at which on the last day they choose the Reina Nacional del Zapallo (the "National Queen of the Pumpkin").[150][151][152] In Portugal the Festival da Abóbora de Lourinhã e Atalaia ("Squashes and Pumpkins Festival in Lourinhã and Atalaia") is held in Lourinhã city, called the Capital Nacional da Abóbora (the "National Capital of Squashes and Pumpkins").[153] Ludwigsburg, Germany annually hosts the world's largest pumpkin festival.[154] In Britain a giant marrow (zucchini) weighing 119 lbs 12oz was displayed in the Harrogate Autumn Flower Show in 2012.[155] In the USA, pumpkin chucking is practiced competitively, with machines such as trebuchets and air cannons designed to throw intact pumpkins as far as possible.[156][157] The Keene Pumpkin Fest is held annually in New Hampshire; in 2013 it held the world record for the most jack-o-lanterns lit in one place, 30,581 on October 19, 2013.[158]

Halloween is widely celebrated with jack-o-lanterns made of large orange pumpkins carved with ghoulish faces, illuminated from inside with candles.[159] The pumpkins used for jack-o-lanterns are C. pepo,[160][161] not to be confused with the ones typically used for pumpkin pie in the United States, which are C. moschata.[162] 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.[163]

See also[edit]

Notes[edit]

  1. ^ 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 genus Cucurbita. Pumpkin and gourd are used to refer to species, varieties, and cultivars commonly referred to by those terms.[4]

References[edit]

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