Cucurbita

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Squash
Squashes at Kew Gardens IncrEdibles 2013.jpg
Cucurbita fruits come in an assortment of colors and sizes.
Cultivated Cucurbita of Canada.jpg
Cross section of a C. pepo pumpkin with the fine-textured flesh favoured for making pumpkin pie
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Cucurbitales
Family: Cucurbitaceae
Tribe: Cucurbiteae
Genus: Cucurbita
L.
Synonyms [1]

Cucurbita ( Latin for ' gourd ') [2] [3] is a genus of herbaceous fruits in the gourd family, Cucurbitaceae (also known as cucurbits or cucurbi), native to the Andes and Mesoamerica. Five edible species are grown and consumed for their flesh and seeds. They are variously known as squash,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 genus Lagenaria , which is in the same family and subfamily as Cucurbita, but in a different tribe, their young fruits are eaten much like those of the Cucurbita species.

Contents

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; they are now cultivated worldwide. 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 annual or short-lived perennial vines which are mesophytic, meaning they require a more or less continuous water supply. The second group are perennials growing in arid zones which are xerophytic, meaning they tolerate dry conditions. Cultivated Cucurbita species were derived from the first group. Growing 5 to 15 meters (15 to 50 feet) in height or length, the plant stem produces tendrils 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 is C. 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, being highly uniform in appearance. [5] The morphological variation in the species C. pepo [6] and C. maxima [7] is so vast that its various subspecies and cultivars have been misidentified as totally separate species. [6]

The typical cultivated Cucurbita species has five-lobed or palmately divided leaves with long petioles, 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 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]

The species are monoecious, with unisexual male (staminate) and female (pistillate) flowers on a single plant 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 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. [8] [11] 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. [8]

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 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 two cotyledons. [11] Fruit size varies considerably: wild fruit specimens can be as small as 4 centimeters (1+12 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 of Switzerland with a 1,054 kg (2,323.7 lb) pumpkin. [14]

Reproductive biology

Cucurbita female flower with pollinating squash bees Peponapis pruinosaCane-12.JPG
Cucurbita female flower with pollinating squash bees

All species of Cucurbita have 20 pairs of chromosomes. [15]

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] [16] [17]

Male flower, part of the perianth removed, arrows indicating nectar pores Cucurbita maxima Zapallo Plomo semilleria Costanzi - flowers detail (masculine) - male flower, some petals and 1 filament removed.jpg
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 called xenia. 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 of C. 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] The plant hormones ethylene and auxin 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] Although Cucurbita 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 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

Kabocha seedling seven days after being sown Kabocha (GH) 21June2005 sown 14June.JPG
Kabocha seedling seven days after being sown

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 12 millimeters (12 in). Seeds planted deeper than 125 millimeters (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. [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 of Cucurbita. A type of stored phosphorus called phytate forms in seed tissues as spherical crystalline intrusions in protein bodies called globoids. Along with other nutrients, phytate is used completely during seedling growth. [33] Heavy metal contamination, including cadmium, 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]

Taxonomy

Cucurbita was formally described in a way that meets the requirements of modern botanical nomenclature by Linnaeus in his Genera Plantarum , [36] the fifth edition of 1754 in conjunction with the 1753 first edition of Species Plantarum . [37] Cucurbita pepo is the type species of the genus. [37] [38] Linnaeus initially included the species C. pepo, C. verrucosa and C. melopepo (both now included in C. pepo), as well as C. citrullus (watermelon, now Citrullus lanatus ) and C. lagenaria (now Lagenaria siceraria ) (both are not Cucurbita but are in the family Cucurbitaceae. [39]

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] [40] The five domesticated species are mostly isolated from each other by sterility barriers and have different physiological characteristics. [40] 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. [41] The buffalo gourd (C. foetidissima) has been used as an intermediary, as it can be crossed with all the common Cucurbita. [11]

An assortment of fruits of C. maxima and C. pepo 2006-10-18Cucurbita pepo06.jpg
An assortment of fruits of C. maxima and C. pepo

Various taxonomic treatments have been proposed for Cucurbita, 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 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: [15] [47]

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]

Distribution and habitat

A festival-winning pumpkin in 2009 weighing 742 kilograms (1,636 lb) 2009 Circleville Pumpkin Show champions.jpg
A festival-winning pumpkin in 2009 weighing 742 kilograms (1,636 lb)

The ancestral species of the genus Cucurbita were present in the Americas before the arrival of humans, [51] [52] and are native to the Americas. 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 the Holocene, whereas the family Cucurbitaceae, represented in Bryonia -like seeds, dates to the Paleocene. [53] Recent genomic studies support the idea that the Cucurbita 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 all Cucurbita species, though the hybrid offspring may not be fertile unless they become polyploid. [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, and C. pepo, originated and were domesticated in Mesoamerica; the fifth, C. maxima, originated and was domesticated in South America. [8]

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. moschata 'Butternut' Cucurbita moschata Butternut 2012 G2.jpg
C. moschata 'Butternut'

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. [8] 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. [8]

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. [8]

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. [5] Types of C. maxima include triloba, [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. Cucurbita moschata 'Tromboncino'.jpg
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 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). [8] 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 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] [64] [65] Debates about the origin of C. 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 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, [67] 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] [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, and cultivars are interfertile. [6] In 1986 Paris proposed a revised taxonomy of the edible cultivated C. pepo based primarily on the shape of the fruit, with eight groups. [57] [69] All but a few C. 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 cultivated C. pepo varieties based on Paris' eight groups and the one non-edible variety
Cultivar group Botanical nameImageDescription
AcornC. 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, [69] ex: Acorn squash [8] [70] [71]
CocozzelleC. pepo var. Ionga Cucurbita pepo Cocozelle fruits.jpg Summer squash, long round slender fruit that is slightly bulbous at the apex, [69] similar to fastigata, ex: Cocozelle von tripolis [8] [70] [71]
CrookneckC. 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, [69] ex: Crookneck squash [8] [70] [71]
PumpkinC. pepo var. pepo
Pumpkin 2 - Evan Swigart.jpg
Winter squash, creeping plant, round, oblate, or oval shape and round or flat on the ends, [69] ex: Pumpkin; [8] [70] [71] includes C. pepo subsp. pepo var. styriaca, used for Styrian pumpkin seed oil [73]
ScallopC. 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, [69] ex: Pattypan squash [8] [70] [71]
StraightneckC. 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, [69] ex: Straightneck squash [8] [70] [71]
Vegetable marrowC. 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, [69] ex: Spaghetti squash (a winter variety) [8] [70] [71]
Zucchini/CourgetteC. 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, [69] similar to fastigata, ex: Zucchini [8] [70] [71]
Ornamental gourdsC. pepo var. ovifera Cucurbita pepo var ovifera (crop).jpg 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 red pumpkin beetle (Aulacophora foveicollis) is a serious pest of cucurbits, especially the pumpkin, which it can defoliate. [78] Cucurbits 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.). [76] Defensive responses to viral, fungal, and bacterial leaf pathogens do not involve cucurbitacin. [79]

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

Cucurbita pepo subsp. texana, from the Grandes Heures of Anne of Brittany, 1503-1508, f. 161, earliest depiction of cucurbits in Europe Bnf Anne f161.jpg
Cucurbita pepo subsp. texana, from the Grandes Heures of Anne of Brittany, 1503–1508, f. 161, earliest depiction of cucurbits in Europe

The genus was part of the culture of almost every native peoples group from southern South America to southern Canada. [52] 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. [50] Species native to North America include C. digitata (calabazilla), [88] and C. foetidissima (buffalo gourd), [89] 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. [90]

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 maize and beans in the region by about 4,000 years. [5] [64] [65] [91] 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. [92] [93] Solid evidence of domesticated C. 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 c. 8,000 years BP the C. pepo peduncles found are consistently more than 10 millimeters (38 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] [94] [95] During the same time frame, average rind thickness increased from 0.84–1.15 millimeters (132364 in). [96] Recent genomic studies suggest that Cucurbita argyrosperma was domesticated in Mexico, in the region that is currently known as the state of Jalisco. [97]

Squash was domesticated first, followed by maize and then beans, becoming part of the Three Sisters agricultural system of companion planting. [98] [99] 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 . [100] Similar words for squash exist in related languages of the Algonquian family. [57] [101]

Production

Squash and gourd* production (2021)
CountryProduction
(millions of tonnes)
Flag of the People's Republic of China.svg  China 7.4
Flag of Ukraine.svg  Ukraine 1.3
Flag of Russia.svg  Russia 1.2
Flag of the United States (23px).png  United States 1.1
Flag of Turkey.svg  Turkey 0.8
Flag of Mexico.svg  Mexico 0.7
Flag of Italy.svg  Italy 0.6
Flag of Indonesia.svg  Indonesia 0.5
Flag of Egypt.svg  Egypt 0.4
World23.4
*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

Cucurbitin is an amino acid and a carboxy pyrrolidine that is found in raw Cucurbita seeds. [103] [104] It retards the development of parasitic flukes when administered to infected host mice, although the effect is seen only if administration begins immediately after infection. [105]

Cucurmosin is a ribosome inactivating protein found in the flesh and seed of Cucurbita, [106] [107] notably Cucurbita moschata .

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. [2] [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, [2] there are occasional reports of cucurbitacin causing illness in humans. [2] Cucurbitacin is also used as a lure in insect traps. [108]

Uses

Nutrition

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
Fat
0.2 g
1.2 g
Vitamins and minerals
Vitamins Quantity
%DV
Vitamin A equiv.
1%
10 μg
1%
120 μg
2125 μg
Thiamine (B1)
4%
0.048 mg
Riboflavin (B2)
11%
0.142 mg
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
9%
262 mg
Zinc
3%
0.29 mg
Other constituentsQuantity
Water95 g

Percentages estimated using US recommendations for adults, [111] except for potassium, which is estimated based on expert recommendation from the National Academies. [112]

As an example of Cucurbita, raw summer squash is 94% water, 3% carbohydrates, and 1% protein, with negligible fat content (table). In a 100-gram reference serving, raw squash supplies 69 kilojoules (16 kcal) of food energy and is rich in vitamin C (20% of the Daily Value, DV), moderate in vitamin B6 and riboflavin (12–17% DV), but otherwise devoid of appreciable nutrient content (table), although the nutrient content of different Curcubita species may vary somewhat. [113]

Pumpkin seeds contain vitamin E, crude protein, B vitamins and several dietary minerals (see nutrition table at pepita). [114] Also present in pumpkin seeds are unsaturated and saturated oils, palmitic, oleic and linoleic fatty acids, [115] as well as carotenoids. [116]

Culinary

The family Cucurbitaceae has many species used as human food. [8] Cucurbita species are some 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 flesh, [109] [117] the fruits and seeds of cultivated varieties are quite edible and need little or no preparation. 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 into pumpkin 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 of C. maxima Sangkhaya fak thong.JPG
Pumpkin custard made from kabocha, a cultivated variant of C. 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 the Pilgrims, who even featured it at the first Thanksgiving. [11] Commercially produced pumpkin commonly used in pumpkin pie is most often varieties of C. moschata; Libby's, by far the largest producer of processed pumpkin, uses a proprietary strain of the Dickinson pumpkin variety of C. 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] The xerophytic 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 a gratin, 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 as cocuzze alla puviredda cooked with olive oil, salt and herbs from Apulia; as torta di zucca from Liguria, or torta di zucca e riso from Emilia-Romagna, the squashes being made into a pie filling with butter, ricotta, parmesan, egg, and milk; and as a sauce for pasta in dishes like spaghetti alle zucchine from Sicily. [129] In Japan, squashes such as small C. moschata pumpkins (kabocha) are eaten boiled with sesame sauce, fried as a tempura dish, or made into balls with sweet potato and Japanese mountain yam. [130]

In culture

Art, music, and literature

Moche squash ceramic. 300 C.E. Larco Museum. Zapallomuseolarco.jpg
Moche squash ceramic. 300 C.E. Larco Museum.

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 in Moche ceramics. [131] [133]

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. [134] [135] 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. [136] 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. [137]

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, [138] and was revived by the Wurzels in Britain on their 2003 album Cutler of the West. [139] [140] John Greenleaf Whittier wrote a poem entitled The Pumpkin in 1850. [141] "The Great Pumpkin" is a fictional holiday figure in the comic strip Peanuts by Charles M. Schulz. [142]

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. [143] [144] Pumpkin is also used in cosmetics. [145]

Folk remedies

Cucurbita have been used in various cultures as folk remedies. Pumpkins have been used by Native Americans to treat intestinal worms and urinary ailments. This Native American remedy was adopted by American doctors in the early nineteenth century as an anthelmintic for the expulsion of worms. [146] In southeastern Europe, seeds of C. pepo were used to treat irritable bladder and benign prostatic hyperplasia. [147] In Germany, pumpkin seed is approved for use by the Commission E, which assesses folk and herbal medicine, for irritated bladder conditions and micturition 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] The FDA 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]

In China, C. moschata seeds were also used in traditional Chinese medicine for the treatment of the parasitic disease schistosomiasis [150] and for the expulsion of tape worms. [151]

In Mexico, herbalists use C. ficifolia in the belief that it reduces blood sugar levels. [152]

Festivals

A pyramid of squashes in the Waterlily House, Kew Gardens, 2013 Squash Pyramid Display.JPG
A 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, Austria, [153] Bolivia, [154] Britain, Canada, [155] Croatia, [156] France, [157] [158] Germany, India, Italy, [159] [160] [161] [162] Japan, [163] Peru, [164] Portugal, Spain, [165] Switzerland, [166] and the United States. Argentina holds an annual nationwide pumpkin festival Fiesta Nacional del Zapallo (lit.'Squashes and Pumpkins National Festival'), in Ceres, Santa Fe, [167] on the last day of which a Reina Nacional del Zapallo (lit.'National Queen of the Pumpkin') is chosen. [168] [169] [170] 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"). [171] Ludwigsburg, Germany annually hosts the world's largest pumpkin festival. [172] In Britain a giant marrow (zucchini) weighing 54.3177 kilograms (119 lb 12 oz) was displayed in the Harrogate Autumn Flower Show in 2012. [173] In the US, pumpkin chucking is practiced competitively, with machines such as trebuchets and air cannons designed to throw intact pumpkins as far as possible. [174] [175] 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. [176]

Hallowe'en is widely celebrated with jack-o-lanterns made of large orange pumpkins carved with ghoulish faces and illuminated from inside with candles. [177] The pumpkins used for jack-o-lanterns are C. pepo, [178] [179] not to be confused with the ones typically used for pumpkin pie in the United States, which are C. moschata. [122] Kew Gardens marked Hallowe’en 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. [180]

See also

Notes

  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]

Related Research Articles

<span class="mw-page-title-main">Gourd</span> Type of fruit

Gourds include the fruits of some flowering plant species in the family Cucurbitaceae, particularly Cucurbita and Lagenaria. The term refers to a number of species and subspecies, many with hard shells, and some without. Many gourds have large, bulbous bodies and long necks, such as Dipper Gourds, many variations of Bottle Gourd and caveman club gourds. One of the earliest domesticated types of plants, subspecies of the bottle gourd, Lagenaria siceraria, have been discovered in archaeological sites dating from as early as 13,000 BC. Gourds have had numerous uses throughout history, including as tools, musical instruments, objects of art, film, and food.

<i>Cucurbita pepo</i> Species of flowering plant that yields varieties of squash and pumpkin

Cucurbita pepo is a cultivated plant of the genus Cucurbita. It yields varieties of winter squash and pumpkin, but the most widespread varieties belong to the subspecies Cucurbita pepo subsp. pepo, called summer squash.

<span class="mw-page-title-main">Kabocha</span> Type of winter squash

Kabocha is a type of winter squash, a Japanese variety of the species Cucurbita maxima. It is also called kabocha squash or Japanese pumpkin in North America. In Japan, "kabocha" may refer to either this squash, to the Western pumpkin, or indeed to other squashes. In Australia, "Japanese pumpkin" is a synonym of Kent pumpkin, a variety of winter squash.

Straightneck squash is a cultivated variety of Cucurbita pepo grown as a type of summer squash that is usually yellow-colored. It is also known as yellow squash, though other squashes, such as crookneck squash, may also be known by that name. It has mildly sweet and watery flesh, and thin tender skins that can be left on the fruit for many types of recipes. It was almost certainly domesticated in the eastern United States, although other variants of the same species were domesticated in Mesoamerica. This squash grows on vined plants reaching 60–90 cm (2.0–3.0 ft) in height that thrive in mild weather. It is well known as an item in American cooking where it is fried, microwaved, steamed, boiled, or baked. It is often used in recipes interchangeably with zucchini. A good yellow summer squash will be small and firm with tender skin free of blemishes and bruising. It is available all year long in some regions, but is at its peak from early through late summer. One similar inedible C. pepo variety is C. pepo var. ovifera.

<span class="mw-page-title-main">Pumpkin seed</span> Seeds of pumpkin and similar squashes

A pumpkin seed, also known as a pepita, is the edible seed of a pumpkin or certain other cultivars of squash. The seeds are typically flat and oval with one axis of symmetry, have a white outer husk, and are light green after the husk is removed. Some pumpkin cultivars are huskless and are grown only for their edible seed. The seeds are nutrient- and calorie-rich, with an especially high content of fat, protein, dietary fiber, and numerous micronutrients. Pumpkin seed can refer either to the hulled kernel or unhulled whole seed and most commonly refers to the roasted end product used as a snack.

<i>Cucurbita ficifolia</i> Plant species cultivated for edible shoots, leaves, flowers, fruit, and seeds

Cucurbita ficifolia is a species of squash, grown for its edible seeds, fruit, and greens. It has common names including black seed squash, chilacayote, cidra, fig-leaf gourd, and Malabar gourd. Compared to other domesticated species in its genus, investigators have noted that samples of C. ficifolia from throughout its range are relatively similar to one other in morphology and genetic composition. Variations do occur in fruit and seed color, some isozymes, and photoperiod sensitivity.

<span class="mw-page-title-main">Eastern Agricultural Complex</span> Agricultural practices of pre-historic native cultures in the eastern United States and Canada

The Eastern Agricultural Complex in the woodlands of eastern North America was one of about 10 independent centers of plant domestication in the pre-historic world. Incipient agriculture dates back to about 5300 BCE. By about 1800 BCE the Native Americans of the woodlands were cultivating several species of food plants, thus beginning a transition from a hunter-gatherer economy to agriculture. After 200 BCE when maize from Mexico was introduced to the Eastern Woodlands, the Native Americans of the eastern United States and adjacent Canada slowly changed from growing local indigenous plants to a maize-based agricultural economy. The cultivation of local indigenous plants other than squash and sunflower declined and was eventually abandoned. The formerly domesticated plants returned to their wild forms.

<i>Cucurbita moschata</i> Species of flowering plant

Cucurbita moschata is a species originating in either Central America or northern South America. It includes cultivars known as squash or pumpkin. C. moschata cultivars are generally more tolerant of hot, humid weather than cultivars of C. maxima or C. pepo. They also generally display a greater resistance to disease and insects, especially to the squash vine borer. Commercially made pumpkin pie mix is most often made from varieties of C. moschata. The ancestral species of the genus Cucurbita were present in the Americas before the arrival of humans. No species within the genus is fully genetically isolated from all the other species. C. moschata can be hybridized with all other species. It has been suggested that this shows that the species of Cucurbita have diversified more recently than those of related genera such as Cucumis and Citrullus.

<span class="mw-page-title-main">Squash bee</span> Bees essential for cucurbit pollination


The name squash bee, also squash and gourd bee, is applied to two related genera of bees in the tribe Eucerini; Peponapis and Xenoglossa. Both genera are oligoleges on the plant genus Cucurbita and closely related plants, although they usually do not visit watermelon, cucumber, and melon plants. They are small genera, containing only 13 and 7 described species, respectively, and their combined range is nearly identical to the range of Cucurbita in the New World, from South America to North America. Their range has become somewhat expanded along with the movement of cucurbits into other areas.

<span class="mw-page-title-main">Crookneck squash</span> Cultivar of Cucurbita pepo

Crookneck squash, also known as yellow squash, is a cultivar of Cucurbita pepo, the species that also includes some pumpkins and most other summer squashes. The plants are bushy and do not spread like the plants of winter squash and pumpkin. Most often used as a summer squash, it is characterized by its yellow skin and sweet yellow flesh, as well as its distinctive curved stem-end or "crooked neck". It should not be confused with crookneck cultivars of Cucurbita moschata, such as the winter squash 'Golden Cushaw', or the vining summer squash 'Tromboncino'. Its name distinguishes it from another similar-looking variety of C. pepo, the straightneck squash, which is also usually yellow. There is one similar non-edible C. pepo variety: C. pepo var. ovifera.

<span class="mw-page-title-main">Winter squash</span> Squash harvested and eaten in mature stage; skin hardened into tough rind

Winter squash is an annual fruit representing several squash species within the genus Cucurbita. Late-growing, less symmetrical, odd-shaped, rough or warty varieties, small to medium in size, but with long-keeping qualities and hard rinds, are usually called winter squash. They differ from summer squash in that they are harvested and eaten in the mature stage when their seeds within have matured fully and their skin has hardened into a tough rind. At this stage, most varieties of this vegetable can be stored for use during the winter. Winter squash is generally cooked before being eaten, and the skin or rind is not usually eaten as it is with summer squash.

<i>Cucurbita maxima</i> Species of squash

Cucurbita maxima, one of at least five species of cultivated squash, is one of the most diverse domesticated species. This species originated in South America from the wild subspecies Cucurbita maxima subsp. andreana over 4,000 years ago. Cucurbita maxima, known for modern varieties as Hubbard, Delicious, Marblehead, Boston Marrow, and Turks Turban, originated in northern Argentina near the Andes or in certain Andean valleys. Secondary centers of diversity include India, Bangladesh, Myanmar, and the southern Appalachians.

<i>Cucurbita digitata</i> Species of vine

Cucurbita digitata is a species of flowering plant in the squash family known by the common names fingerleaf gourd and bitter squash. It is similar to Cucurbita californica, Cucurbita cordata, Cucurbita cylindrata, and Cucurbita palmata and all these species hybridize readily. These species form the only restricted xerophyte species group in the genus Cucurbita. Each member of this species group is native to the Southwestern United States and Northwestern Mexico where they are relatively uncommon. Each group member is found in hot, arid regions with low rainfall. They prefer soil that is loose, gravelly, and well-drained. C. digitata is native to northern Baja California at higher elevations, northern Sonora, Mexico, southern Arizona, and southwestern New Mexico. The juvenile leaves of C. cylindrata, C. cordata, C. digitata, and C. palmata show a high degree of similarity, but their mature leaves are visibly different, as are their root structures. C. palmata and C. digitata are sympatric, with C. palmata separating the ranges of C. digitata at the juncture of Baja California, California, and Arizona. C. digitata fruits are clear green mottle that turns yellow at maturity, striped, and round.

<span class="mw-page-title-main">Pumpkin</span> Category of culinary winter Cucurbita squashes

A pumpkin is a cultivated winter squash in the genus Cucurbita. The term is most commonly applied to round, orange-colored squash varieties, but does not possess a scientific definition. It may be used in reference to many different squashes of varied appearance and belonging to multiple species in the Cucurbita genus.

<i>Cucurbita argyrosperma</i> Species of plant

Cucurbita argyrosperma, also called the cushaw squash and silver-seed gourd, is a species of winter squash originally from the south of Mexico. This annual herbaceous plant is cultivated in the Americas for its nutritional value: its flowers, shoots, and fruits are all harvested, but it is cultivated most of all for its seeds, which are used for sauces. It was formerly known as Cucurbita mixta.

<i>Cucurbita okeechobeensis</i> Species of vine

Cucurbita okeechobeensis, the Okeechobee gourd, is a species of gourd in the family Cucurbitaceae, native to Mexico and the United States. There are two subspecies; one is endemic to Florida, primarily in the region around Lake Okeechobee, the other to the State of Veracruz in eastern Mexico. Once abundant, it has state and federal listing as an endangered species. One of its peculiarities is the yellow corolla not so common in other Cucurbita species.

<i>Cucurbita ecuadorensis</i> Species of flowering plant

Cucurbita ecuadorensis is a species of squash, described in 1965 as growing wild in Ecuador. Like most wild gourds and squashes, it is a creeping vine and is often found climbing over other vegetation. It has been found only in the western provinces of Guayas and Manabí. There is evidence that it was domesticated in Ecuador around 10,000 years ago, likely for its seeds, but no direct records exist and it is no longer cultivated. It is resistant to many diseases of cultivated Cucurbita species, and has been used to breed resistance to several diseases into common squashes. For example, researchers at Cornell University used Cucurbita ecuadorensis to breed resistance to papaya ringspot virus, watermelon mosaic virus, and powdery mildew, into common Cucurbita maxima cultivars. Cucurbita ecuadorensis is listed on the IUCN Red List as vulnerable and is found protected in the Machalilla National Park.

Cucurbita lundelliana is a mesophyte plant species of the genus Cucurbita. It is native to Mexico, Guatemala, and Belize. It has not been domesticated. It is found in the Yucatán region near sea level among limestone cliffs. In Guatemala it is found in Parque Nacional Yaxha Nakum Naranjo along Rio Ixtinto and near Laguna Julequito.

Cucurbita californica is a species of flowering plant in the squash family.

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