From Wikipedia, the free encyclopedia - View original article

Cucurbita 2011 G1.jpg
Cucurbita fruits come in an assortment of colors and sizes
Scientific classification

see below

Jump to: navigation, search
Cucurbita 2011 G1.jpg
Cucurbita fruits come in an assortment of colors and sizes
Scientific classification

see below


Cucurbita (Latin for gourd)[2] is a genus in the gourd family Cucurbitaceae native to and originally cultivated in the Andes and Mesoamerica. The Cucurbita genus is an important source of human food and is used for other purposes such as beverages, medicine, oil, and 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, are grown for their edible fruits and seeds. Real (bottle-)gourds, 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 different tribes.

There are five domesticated species: Cucurbita argyrosperma, C. ficifolia, C. maxima, C. moschata, and C. pepo. C. pepo includes varieties of both winter squash and summer squash, and C. moschata can be used as winter squash because the full-grown fruits can be stored for months. There is no universal agreement on how to handle the taxonomic treatment of the genus, with the number of species being listed varying from 13–30. There is uncertainty as to the ancestors of some species.

Most Cucurbita species are vines that grow several meters in length, have yellow or orange flowers, and may have tendrils. Non-vining cultivars of C. pepo have been developed. There are two types of flowers on a 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 pollinators in the apid group, but other insects including honey bees also visit.

The Cucurbita genus is an important source of human food and the fruits are good sources of several nutrients such as vitamin A, vitamin C, dietary fiber, niacin, folic acid, and iron. In addition, they are free of fat and cholesterol. The plants also contain the toxins cucurmosin and cucurbitacin. Medical uses of the plant include treating skin conditions and improving visual acuity.


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

Most Cucurbita species are climbing annual vines and are mesophytes (requiring adequate water), while the perennials grow in tropical zones and are xerophytes (tolerating dry conditions). The plant stem can grow 5 to 15 meters (16 to 49 ft) and produces tendrils that help it climb adjacent plants and structures or along the ground. Most species do not readily root from the nodes; Cucurbita ficifolia and the 4 cultivated mesophytes are a notable exception. 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, even within a single species. C. ficifolia is an exception to this, being highly uniform in appearance.[3] The morphological differences in the species C. pepo[4] and C. maxima[5] 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 (radiating from the base) leaves having long petioles; with 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, and sometimes these are hardened and sharp. Tendrils grow from each node, and are spring-like; in some species the tendrils may branch. 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.[6][7] Female flowers have thick pedicels, and an inferior ovary with 3–5 stigmas that each have two lobes.[8][9] C. argyrosperma has ovate-cordate leaves and the corollas of its female flowers are larger than the male flowers. The shape of C. pepo leaves varies widely; its female flowers have a small calyx. C. moschata plants can have light or dense pubescence. The calyx of its male flowers is comparatively short. C. ficifolia leaves are slightly angular and have light pubescence. Its female flowers have noticeably larger corollas than the male flowers.[9] The leaves of all four of these species may or may not have white spots on their leaves.[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 formed from hypanthium tissue fused to the exocarp. The fleshy interior is 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 considered pepos.[10][11][12] Cucurbita fruits are large and fleshy.[6] 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.[8] 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).[9] The current world record was set by Ron Wallace of Greene, Rhode Island with a 911.2 kilograms (2,009 lb) Atlantic Giant pumpkin.[13]

Reproductive biology[edit]

Cucurbita female flower with pollinating squash bees

Many North and Central American species are visited by specialist pollinators in the apid group Eucerini, especially the genera Peponapis and Xenoglossa, and these squash bees can be crucial to the flowers producing fruit after pollination.[3][14] 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.[15] Parthenocarpy is known to occur in certain cultivars of C. pepo.[16][17]

Male flower, part of the perianth and 1 filament removed

The plant hormone ethylene promotes the production of female flowers. When a plant already has fruit, new female flowers are inhibited, and male flowers are more frequent, an effect that appears to be due to reduced natural ethylene production within the plant stem.[18] 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] Ethylene and the hormone auxin are key in fruit set and development.[20] The most critical factors in flowering and fruit set are physiological, not climatic, factors.[21] If a fruit is developing, then subsequent female flowers on the plant are less likely to mature, a phenomenon called "first-fruit dominance".[21]

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.[22] Gibberellin is also involved in other developmental processes of plants such as seed and stem growth.[23]

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. Maximum seed germination in C. moschata occurs 45 days after anthesis. Seed weight is at its maximum 75 days after germination.[24] 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.[25] C. foetidissima seeds have a 90% germination rate in pH 8 soil. Plants younger than 19 days old are not able to sprout from the roots after removing the shoots. Over 90% of plants sprout after 29 days from planting.[26]

High levels of pollen load are known to produce more seeds in fruits, faster and more likely seed germination, larger fruits, greater likelihood of fruit maturation, more plant biomass, and larger stems.[27] 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 into spherical crystalline intrusions in protein bodies called globoids. The nutrients in globoids are eventually used completely during seedling growth.[28] Heavy metal contamination, including cadmium, has a significant negative impact on plant growth.[29] Sterols are critical to the plant life cycle of Cucurbita and stigmastadienol is the most critical of them. More study is needed on sterol biosynthesis.[30] Sterol levels fluctuate during the plant life cycle, such as decreasing at the onset of flowering. Cucurbita plants grown in the spring tend to grow larger than those grown in the autumn.[31]

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,[32][33] and all species are native to the New World since the early development of agriculture. The likely center of origin is southern Mexico and then spreading south into South America, largely in what is now known as Mesoamerica, and north to what is now the southwestern United States.[32] Evolutionarily speaking, the genus is relatively recent in origin, and no species within the genus is genetically isolated from all the other species. Cucurbita moschata acts as the genetic bridge within the genus.[34] The genus was part of the culture of almost every native peoples group from southern South America to southern Canada.[33] Modern-day cultivated Cucurbita are not found in the wild.[3] 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.[35] Species native to North America include C. digitata (calabazilla),[36] and C. foetidissima (buffalo gourd),[37] C. palmata (coyote melon), and C. pepo.[3] Every species of Cucurbita has 20 pairs of chromosomes.[38] Some species, such as C. digitata and C. ficifolia, are referred to as gourds, but the real (bottle-)gourds, 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 different tribes.[39]

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.[3][40][41][42] 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.[43][44] 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.[8][45][46] During the same time frame, average rind thickness increased from 0.84 millimeters (0.033 in) to 1.15 millimeters (0.045 in).[47]

Moche Squash Ceramic. 300 A.D. Larco Museum

The process of domesticating the squashes 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.[48][49] All three crops have been frequently depicted in the art work of the native peoples of the Americas for at least 2,000 years.[50][51] For example, squashes are represented frequently in Moche ceramics.[50][52]

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.[53] Similar words for squash exist in related languages of the Algonquian family.[54][55]

Cucurbita began to spread to other parts of the world after Christopher Columbus's arrival in the New World in 1492.[56][57] Until 1992, the earliest known depictions of this genus in Europe was of Cucurbita pepo in De Historia Stirpium Commentarii Insignes in 1542 by Leonhart Fuchs, a German botanist. In 1992, two paintings, one of C. pepo and one of C. maxima, were found in a festoon at Villa Farnesina in Rome.[58] These were painted between 1515 and 1518. In 2001 depictions of this genus were found in Grandes Heures of Anne of Brittany (Les Grandes Heures d'Anne de Bretagne), a French devotional book printed 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.[59]


The full phylogeny of this genus is unknown, and research is ongoing in 2014.[60][61] The following cladogram of Cucurbita phylogeny is based upon a 2002 study based on mitochondrial DNA by Sanjur et al.[35]

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


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.[62][63] 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.[3][34] The five domesticated species are mostly isolated from each other by sterility barriers and have different physiological characteristics.[34] 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.[64] The buffalo gourd (C. foetidissima), which does not taste good, has been used as an intermediary as it can cross breed with all the common Cucurbita.[8]

Various taxonomic treatments have been proposed for Cucurbita, ranging from 13–30 species.[2] In 1990, Michael Nee classified them into the following 13 species groups (27 species total), listed by group and alphabetically, with geographic origin:[3][65][66]

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:[38][69]

Habitat and distribution[edit]

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

Of the five domesticated species, four of them: Cucurbita argyrosperma, C. ficifolia, C. moschata, and C. pepo originated and were domesticated in Mesoamerica; whereas in the case of C. maxima this occurred in South America. The exact location of origin of C. ficifolia and C. moschata is less certain.[9]

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.[55] In a 1989 study on the origins and development of C. pepo, 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.[55]

Cucurbita argyrosperma is not as widespread as the other species but is found from Mexico to Nicaragua. It has large seeds that are high in oil and protein and was likely 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–June and fruit harvested from October–December.[9]

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,[35] most likely Argentina and Uruguay. They are sensitive to frost, prefer sunlight, and thrive in soil with a pH of 6.0 to 7.0.[70] It did not start to spread into North America until after the arrival of Columbus. By the 16th century varieties were in use by native peoples of the United States.[3] Groups of C. maxima include Triloba,[71] Zapallito,[72] Zipinka,[73] Banana, Delicious, Hubbard, Marrow (C. maxima Marrow), Show, and Turban.[74]

C. moschata 'Butternut'

Cucurbita moschata is native to Latin America but the precise location of origin is uncertain.[75] 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).[9] Groups of C. moschata include: Cheese, Crookneck (C. moschata), and Bell.[74]

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.[3][9][40][41] Debates about the origin of C. pepo have been going on since at least 1857.[76] 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.[3] A more recent theory by botanist Andres in 1987 is that it is a descendant of C. fraterna and hybridized with C. texana;[77] 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.[9][35][77][78] C. pepo may have appeared in the Old World prior to moving from Mexico into South America.[9] 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.[4] In 1986 botanist Paris proposed a taxonomy of C. pepo consisting of eight edible groups based on their basic shape.[55][79] All but a few C. pepo can be included in these groups.[79] These eight edible cultivated varieties of C. pepo vary widely in shape and color,[9][80][81] and one non-edible cultivated variety:[82]

Cultivar groupBotanical nameImageDescription
AcornC. pepo var. turbinataStarr 070730-7820 Cucurbita pepo.jpgwinter squash, both a shrubby and creeping plant, obovoid or conical shape, pointed at the apex and with longitudinal grooves, thus resembling a spinning top,[79] ex: Acorn squash[9][80][81]
CocozzelleC. pepo var. IongaCucurbita pepo Cocozelle fruits.jpgsummer squash, long round slender fruit that is slightly bulbous at the apex,[79] similar to fastigata, ex: Cocozelle von tripolis[9][80][81]
CrookneckC. pepo var. torticollia (also torticollis)Crooked Neck Squash.jpgsummer 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,[79] ex: Yellow crookneck squash[9][80][81]
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,[79] ex: Pumpkin;[9][80][81] includes C. pepo subsp. pepo var. styriaca, used for Styrian pumpkin seed oil[83]
ScallopC. pepo var. clypeata; called C. melopepo by Linnaeus[4]Pattypan squash J1.jpgsummer squash, prefers half-shrubby habitat, flattened or slightly discoidal shape, with undulations or equatorial edges,[79] ex: Pattypan squash[9][80][81]
StraightneckC. pepo var. recticollisCucurbita pepo var recticollis, Yellow Straightneck Squash (crop).jpgsummer squash, shrubby plant, with yellow or golden fruit and verrucose rind, similar to var. torticollia but a stem end that narrows,[79] ex: Yellow summer squash[9][80][81]
Vegetable marrowC. pepo var. fastigataSpaghetti Squash 700.jpgsummer and winter squashes, creeper traits and a semi-shrub, cream to dark green color, short round fruit with a slightly broad apex,[79] ex: Spaghetti squash (a winter variety)[9][80][81]
ZucchiniC. pepo var. cylindricaZucchini-Whole.jpgsummer squash, presently the most common group of cultivars, origin is recent (19th century), semi-shrubby, cylindrical fruit with a mostly consistent diameter,[79] similar to fastigata, ex: Zucchini[9][80][81]
Ornamental gourdsC. pepo var. oviferaCucurbita pepo var ovifera (crop).jpgnon-edible,[82] field pumpkins 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.[40]

Culinary uses[edit]

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

The species of Cucurbita are prepared and consumed in many ways. While the skins are often bitter, the fruits and seeds of cultivated varieties are quite edible by humans with little or no preparation. The seeds and fruits of most varieties can be stored for long periods of time.[3] In addition, the young leaves, young shoot tips, and flowers can be consumed.[84] The Cucurbitaceae family has the largest number of different species used as human food of any plant family.[9] Cucurbita is one of the most important of those, either for their fruit or the seeds within. The winter varieties have thick, inedible skins, and so store well. They have a sweet taste. In contrast, summer squash have a thin, edible skin. The seeds of both types can be ground into a flour or meal,[85] roasted, eaten raw, made into pumpkin seed oil,[83] or otherwise prepared. Buffalo gourd oil is made from Cucurbita foetidissima.[86] By the time of European contact Cucurbita had long been a major food source for the native peoples of the Americas. They became an important food for the European settlers, including the Pilgrims, for whom they were a major part of the first Thanksgiving.[8] The large red-orange squashes often seen at Halloween in the United States are C. maxima, but not to be confused with the orange type used for jack-o-lanterns, which are C. pepo.[3] 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 pumpkins.[87] 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,[88] salads, soups, and stuffing.[89] The species that are xerophytes are proving useful in the search for nutritious foods that grow well in arid regions.[90]

Medicinal uses[edit]

Cucurbita is used in cosmetics for dry and sensitive skin and in treating schistosomiasis.[91][92][93] The flesh of Cucurbita argyrosperma is used for treating burns and skin conditions while its seeds are treated with water and used as an anesthetic and to promote lactation in nursing women.[9] C. ficifolia is used to make flavorings for soft and mildly alcoholic drinks,[9] and high doses of C. ficifolia have been shown to be successful in reducing blood sugar levels.[2] The fruits of some species, such as C. foetidissima, are used as soaps and detergents.[2][86] Cucurbita fruits are an important source for humans of carotenoids, vitamin A, and rhodopsin, all of which are important to good visual acuity.[8][94] Cucurbitin is an amino acid and a carboxypyrrolidine found in Cucurbita seeds that can eliminate parasitic worms.[95][96]


Summer squash, all varieties, raw
Nutritional value per 100 g (3.5 oz)
Energy69 kJ (16 kcal)
3.4 g
Sugars2.2 g
Dietary fiber1.1 g
0.2 g
1.2 g
Vitamin A equiv.
10 μg
120 μg
2125 μg
Thiamine (B1)
0.048 mg
Riboflavin (B2)
0.142 mg
Niacin (B3)
0.487 mg
0.155 mg
Vitamin B6
0.218 mg
Folate (B9)
29 μg
Vitamin C
17 mg
Vitamin K
3 μg
Trace metals
0.35 mg
17 mg
0.175 mg
38 mg
262 mg
0.29 mg
Other constituents
Water95 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. Pumpkins have anti-diabetic, antioxidant, anticarcinogen, and anti-inflammatory pharmacological properties include.[97] Pumpkins and pumpkin seeds have high levels of crude protein, calcium, iron, potassium, phosphorus, magnesium, zinc,[98][99] and beta-carotene.[100] Squashes are good sources of vitamin A, vitamin C, potassium, dietary fiber, niacin, folic acid, and iron. They are free of fat and cholesterol.[101]


Cucurmosin is a ribosome inactivating protein found in the sarcocarp (flesh) and seed of Cucurbita,[102][103] notably Cucurbita moschata. Cucurmosin is more toxic to cancer cells than healthy cells.[102][104]

Cucurbitacin is a plant steroid poisonous to mammals found in wild Cucurbita in quantities sufficient to discourage herbivores.[105] It has been bred out of cultivated varieties. It has a bitter taste and ingesting too much can cause stomach cramps, diarrhea and even collapse.[106] While the process of domestication has largely removed the bitterness from the cultivated varieties,[2] there are occasional reports of cucurbitacin getting into the human food supply and causing illnesses.[2] This is what makes wild Cucurbita and most ornamental gourds, with the exception of an occasional C. fraterna and C. sororia, bitter to taste.[2][77]

Pests and diseases[edit]

Cucurbita species are used as food plants by the larvae of some Lepidoptera species, including Cabbage Moths, Hypercompe indecisa, and Turnip Moths.[95] Cucurbita can be susceptible to the pest Bemisia argentifolii (silverleaf whitefly)[107] 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).[108] The squash bug causes major damage to plants because of its very toxic saliva.[109] 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.).[108] Defensive responses to viral, fungal, and bacterial leaf pathogens do not involve cucurbitacin.[105]

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),[110] Watermelon mosaic virus (WMV), and Zucchini yellow mosaic virus (ZYMV).[111][112][113][114] PRSV is the only one of these viruses that does not affect all cucurbits, which is the family Cucurbitaceae.[111][115] SqMV and CMV are the most common viruses among cucurbits.[116][117] Symptoms of these viruses show a high degree of similarity and laboratory investigation is often needed to differentiate which one is affecting plants.[110]


Squashes are primarily grown for the fresh food market.[118] 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:[119]

Top ten squash producers — 2012[119]
(metric tons)
China China6,140,840
India India4,424,200
Russia Russia988,180
United States USA778,630
Iran Iran695,600
Egypt Egypt658,234
Mexico Mexico522,388
Ukraine Ukraine516,900
Italy Italy508,075
Turkey Turkey430,402
Top 10 total15,663,449

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.[119] 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.[118]

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

Top ten cucumber producers — 2012[119]
(metric tons)
China China40,710,200
Iran Iran1,811,630
Turkey Turkey1,739,190
Russia Russia1,161,870
United States USA883,360
Ukraine Ukraine860,100
Spain Spain682,900
Egypt Egypt631,408
Japan Japan587,800
Indonesia Indonesia547,141
Top 10 total49,075,599
Top ten watermelon producers — 2012[119]
(metric tons)
China China56,649,725
Turkey Turkey3,683,100
Iran Iran3,466,880
Brazil Brazil1,870,400
United States USA1,866,660
Egypt Egypt1,637,090
Uzbekistan Uzbekistan1,182,400
Russia Russia1,151,580
Mexico Mexico1,036,800
Algeria Algeria946,200
Top 10 total73,490,835

See also[edit]


  1. ^ "Cucurbita L.". Tropicos, Missouri Botanical Garden. Retrieved January 12, 2014. 
  2. ^ a b c d e f g Burrows, George E.; Tyrl, Ronald J. (2013). Toxic Plants of North America. Oxford: Wiley-Blackwell. pp. 389–391. ISBN 978-0-8138-2034-7. 
  3. ^ a b c d e f g h i j k l Nee, Michael (1990). "The Domestication of Cucurbita (Cucurbitaceae)". Economic Botany (New York: New York Botanical Gardens Press) 44 (3, Supplement: New Perspectives on the Origin and Evolution of New World Domesticated Plants): 56–68. doi:10.2307/4255271. JSTOR 4255271.  edit
  4. ^ a b c Decker-Walters, Deena S.; Staub, Jack E.; Chung, Sang-Min; Nakata, Eijiro; Quemada, Hector D. (2002). "Diversity in Free-Living Populations of Cucurbita pepo (Cucurbitaceae) as Assessed by Random Amplified Polymorphic DNA". Systematic Botany (American Society of Plant Taxonomists) 27 (1): 19–28. doi:10.2307/3093892. JSTOR 3093892.  edit
  5. ^ Millán R, 1945. Variaciones del zapallito amargo Cucurbita andreana y el origen de Cucurbita maxima. Revista Argentina de Agronomía 12:86-93. And literature cited.
  6. ^ a b Mabberley, D. J. (2008). The Plant Book: A Portable Dictionary of the Vascular Plants. Cambridge: Cambridge University Press. p. 235. ISBN 978-0-521-82071-4. 
  7. ^ Lu, Anmin; Jeffrey, Charles. "Cucurbita L.". Flora of China. pp. 55–56. 
  8. ^ a b c d e f "Cucurbitaceae--Fruits for Peons, Pilgrims, and Pharaohs". University of California at Los Angeles. Retrieved September 2, 2013. 
  9. ^ a b c d e f g h i j k l m n o p q r s t u v Saade, R. Lira; Hernández, S. Montes. "Cucurbits". Purdue Horticulture. Retrieved September 2, 2013. 
  10. ^ "A Systematic Treatment of Fruit Types". Worldbotanical. Retrieved October 10, 2013. 
  11. ^ "Pepo". Royal Botanical Gardens, Kew. Retrieved October 10, 2013. 
  12. ^ Schrager, Victor (2004). The Compleat Squash: A Passionate Grower's Guide to Pumpkins, Squash, and Gourds. New York: Artisan. p. 25. ISBN 978-1-579-65251-7. 
  13. ^ Marturana, Amy (October 5, 2012). "Ron Wallace Grows The Largest Pumpkin, Breaking World Record". Huffington Post. Retrieved October 29, 2013. 
  14. ^ Hurd, Paul D.; Linsley, E. Gorton (1971). "Squash and Gourd Bees (Peponapis, Xenoglossa) and the Origin of the Cultivated Cucurbita". Evolution (St. Louis, MO: Society for the Study of Evolution) 25 (1): 218–234. doi:10.2307/2406514. JSTOR 2406514.  edit
  15. ^ Stephenson, Andrew G.; Devlin, B.; Horton, J. Brian (1988). "The Effects of Seed Number and Prior Fruit Dominance on the Pattern of Fruit Production in Cucurbits pepo (Zucchini Squash)". Annals of Botany (Oxford Journals) 62 (6): 653–661. 
  16. ^ Robinson, R. W.; Reiners, Stephen (July 1999). "Parthenocarpy in Summer Squash" (PDF). HortScience 34 (4): 715–717. 
  17. ^ Menezes, C. B.; Maluf, W. R.; Azevedo, S. M.; Faria, M. V.; Nascimento, I. R.; Gomez, L. A.; Bearzoti, E. (March 2005). "Inheritance of Parthenocarpy in Summer Squash (Cucurbita pepo L.).". Genetics and Molecular Research 31 (4): 39–46. PMID 15841434. 
  18. ^ Krupnick, Gary A.; Brown, Kathleen M.; Stephenson, Andrew G. (1999). "The Influence of Fruit on the Regulation of Internal Ethylene Concentrations and Sex Expression in Cucurbita texana". International Journal of Plant Sciences 160 (2): 321–330. JSTOR 10.1086/314120. 
  19. ^ Murray, M. (1987). "Field Applications Of Ethephon For Hybrid And Open-Pollinated Squash (Cucurbita Spp) Seed Production". Acta Horticulturae 201: 149–156. 
  20. ^ Martínez, Cecelia; Manzano, Susana; Megías, Zoraida; Garrido, Dolores; Picó, Belén; Jamilena, Manuel (2013). "Involvement of Ethylene Biosynthesis and Signalling in Fruit Set and Early Fruit Development in Zucchini Squash (Cucurbita pepo L.)". BMC Plant Biology 13 (139). doi:10.1186/1471-2229-13-139. 
  21. ^ a b Stapleton, Suzanne Cady; Wien, H. Chris; Morse, Roger A. (2000). "Flowering and Fruit Set of Pumpkin Cultivars under Field Conditions" (PDF). HortScience 35 (6): 1074–1077. 
  22. ^ Pimenta Lange, Maria João; Knop, Nicole; Lange, Theo (2012). "Stamen-derived Bioactive Gibberellin is Essential for Male Flower Development of Cucurbita maxima L.". Journal of Experimental Botany 63 (7): 2681–2691. doi:10.1093/jxb/err448. PMC 3346225. PMID 22268154. 
  23. ^ "Plant Hormones". Charles Sturt University. Retrieved January 15, 2014. 
  24. ^ Wilson, Mack A.; Splittstoesser, Walter E. (1980). "The Relationship Between Embryo Axis Weight and Reserve Protein During Development and Pumpkin Seed Germination". Journal of Seed Technology 5 (2): 35–41. JSTOR 23432821.  edit
  25. ^ Oliver, Lawrence R.; Harrison, Steve A.; McClelland, Marilyn (1983). "Germination of Texas Gourd (Cucurbita texana) and Its Control in Soybeans (Glycine max)". Weed Science 31 (5): 700–706. JSTOR 4043694.  edit
  26. ^ Horak, Michael J.; Sweat, Jonathan K. (1994). "Germination, Emergence, and Seedling Establishment of Buffalo Gourd (Cucurbita foetidissima)". Weed Science 42 (3): 358–363. JSTOR 4045510.  edit
  27. ^ Winsor, J. A.; Davis, L. E.; Stephenson, A. G. (1987). "The Relationship Between Pollen Load and Fruit Maturation and the Effect of Pollen Load on Offspring Vigor in Cucurbita pepo". The American Naturalist 129 (5): 643–656. JSTOR 2461727.  edit
  28. ^ Beecroft, Penny; Lott, John N. A. (1996). "Changes in the Element Composition of Globoids From Cucurbita maxima and Cucurbita andreana Cotyledons During Early Seedling Growth". Canadian Journal of Botany 74 (6): 838–847. doi:10.1139/b96-104. 
  29. ^ Subin, M. P.; Francis, Steffy (2013). "Phytotoxic Effects of Cadmium on Seed Germination, Early Seedling Growth and Antioxidant Enzyme Activities in Cucurbita maxima Duchesne". International Research Journal of Biological Sciences 2 (9): 40–47. doi:10.1139/b96-104. 
  30. ^ Fenner, G. P.; Raphiou, I. (1995). "Growth of Cucurbita maxima L. Plants in the Presence of the Cycloartenol Synthase Inhibitor U18666A". Lipids 30 (3): 253–256. doi:10.1007/bf02537829. PMID 7791534. 
  31. ^ Fenner, G. P.; Patteron, G. W.; Lusby, W. R. (1989). "Developmental Regulation of Sterol Biosynthesis in Cucurbita maxima L.". Lipids 24 (4): 271–277. doi:10.1007/BF02535162. 
  32. ^ a b Bemis, W. P.; Whitaker, Thomas W. (April 1969). "The Xerophytic Cucurbita of Northwestern Mexico and Southwestern United States". Madroño (California Botanical Society) 20 (2): 33–41. JSTOR 41423342.  edit
  33. ^ a b Smith, Bruce D. (15 August 2006). "Eastern North America as an Independent Center of Plant Domestication". Proceedings of the National Academy of Sciences of the United States of America (National Academy of Sciences of the United States of America) 103 (33): 12223–12228. doi:10.1073/pnas.0604335103. PMC 1567861. PMID 16894156. 
  34. ^ a b c Whitaker, T.W.; Bemis, W.P. (1975). "Origin and Evolution of the Cultivated Cucurbita". Bulletin of the Torrey Botanical Club 102 (6): 362–368. JSTOR 2484762. 
  35. ^ a b c d Sanjur, Oris I.; Piperno, Dolores R.; Andres, Thomas C.; Wessel-Beaver, Linda (2002). "Phylogenetic Relationships among Domesticated and Wild Species of Cucurbita (Cucurbitaceae) Inferred from a Mitochondrial Gene: Implications for Crop Plant Evolution and Areas of Origin" (PDF). Proceedings of the National Academy of Sciences of the United States of America (Washington, DC: National Academy of Sciences) 99 (1): 535–540. doi:10.1073/pnas.012577299. JSTOR 3057572.  edit
  36. ^ "Cucurbita digitata A. Gray". United States Department of Agriculture. Retrieved September 1, 2013. 
  37. ^ "Cucurbita ficifolia Bouché". United States Department of Agriculture. Retrieved September 1, 2013. 
  38. ^ a b Rhodes, A. M.; Bemis, W. P.; Whitaker, Thomas W.; Carmer, S. G. (1968). "A Numerical Taxonomic Study of Cucurbita". Brittonia (New York Botanical Garden Press) 20 (3): 251–266. doi:10.2307/2805450. JSTOR 2805450.  edit
  39. ^ Roberts, Katherine M. "Cucurbitaceae". Washington University in St. Louis. Retrieved September 1, 2013. 
  40. ^ a b c Wilson, Hugh D. "What is Cucurbita texana?". Free-living Cucurbita pepo in the United States Viral Resistance, Gene Flow, and Risk Assessment. Texas A&M Bioinformatics Working Group. Retrieved September 8, 2013. 
  41. ^ a b Gibbon, Guy E.; Ames, Kenneth M. (1998). Archaeology of Prehistoric Native America: An Encyclopedia. New York: Routledge. p. 238. ISBN 978-0-815-30725-9. 
  42. ^ Roush, Wade (9 May 1997). "Archaeobiology: Squash Seeds Yield New View of Early American Farming". Science (American Association For the Advancement of Science) 276 (5314): 894–895. doi:10.1126/science.276.5314.894. 
  43. ^ Schoenwetter, James (April 1974). "Pollen Records of Guila Naquitz Cave". American Antiquity (Society for American Archaeology) 39 (2): 292–303. doi:10.2307/279589. JSTOR 279589.  edit
  44. ^ Benz, Bruce F. (2005). "Archaeological evidence of teosinte domestication from Guilá Naquitz, Oaxaca". Proceedings of the National Academy of Sciences (National Academy of Sciences of the United States of America) 98 (4): 2104–2106. doi:10.1073/pnas.98.4.2104. 
  45. ^ Smith, Bruce D. (22 December 1989). "Origins of Agriculture in Eastern North America". Science (Washington, DC: American Association for the Advancement of Science) 246 (4937): 1566–1571. doi:10.1126/science.246.4937.1566. PMID 17834420. 
  46. ^ Smith, Bruce D. (May 1997). "The Initial Domestication of Cucurbita pepo in the Americas 10,000 Years Ago". Science (Washington, DC: American Association for the Advancement of Science). doi:10.1126/science.276.5314.932. 
  47. ^ Feinman, Gary M.; Manzanilla, Linda (2000). Cultural Evolution: Contemporary Viewpoints. New York: Kluwer Academic. p. 31. ISBN 978-0-306-46240-5. 
  48. ^ Landon, Amanda J. (2008). "The "How" of the Three Sisters: The Origins of Agriculture in Mesoamerica and the Human Niche". Nebraska Anthropologist (Lincoln, NE: University of Nebraska-Lincoln): 110–124. 
  49. ^ Bushnell, G. H. S. (1976). "The Beginning and Growth of Agriculture in Mexico". Philosophical Transactions of the Royal Society of London (London: Royal Society of London) 275 (936): 117–120. doi:10.1098/rstb.1976.0074. 
  50. ^ a b "Moche Decorated Ceramics". Metropolitan Museum of Art. Retrieved October 10, 2013. 
  51. ^ Benson, Elizabeth P. (1983). "A Moche "Spatula"". Metropolitan Museum Journal (The University of Chicago Press) 18: 39–52. JSTOR 1512797.  edit
  52. ^ Berrin Larco Museum, Katherine (1997). The Spirit of Ancient Peru: Treasures from the Museo Arqueológico Rafael Larco Herrera. New York: Thames and Hudson. ISBN 978-0-500-01802-6. 
  53. ^ "How Did the Squash Get its Name?". Library of Congress. Retrieved October 10, 2013. 
  54. ^ Cutler, Charles L. (2000). O Brave New Words: Native American Loanwords in Current English. Norman, OK: University of Oklahoma Press. pp. 39–42. ISBN 978-0-8061-3246-4. 
  55. ^ a b c d Paris, Harry S. (1989). "Historical Records, Origins, and Development of the Edible Cultivar Groups of Cucurbita pepo (Cucurbitaceae)". Economic Botany (New York Botanical Garden Press) 43 (4): 423–443. JSTOR 4255187.  edit
  56. ^ Whitaker, Thomas W. (1947). "American Origin of Cultivated Cucurbits". Annals of the Missouri Botanical Garden (St. Louis, MO: Missouri Botanical Garden Press) 34: 101–111. doi:10.2307/2394459. JSTOR 2394459.  edit
  57. ^ Whitaker, Thomas W. (1956). "The Origin of the Cultivated Cucurbita". The American Naturalist (Chicago, IL: University of Chicago Press) 90 (852): 171–176. doi:10.1086/281923. JSTOR 2458406.  edit
  58. ^ Janick, Jules; Paris, Harry S. (February 2006). "The Cucurbit Images (1515–1518) of the Villa Farnesina, Rome". Annals of Botany (Oxford) 97 (2): 165–176. doi:10.1093/aob/mcj025. PMC 2803371. PMID 16314340. 
  59. ^ Paris, Harry S.; Daunay, Marie-Christine; Pitrat, Michel; Janick, Jules (July 2006). "First Known Image of Cucurbita in Europe, 1503–1508". Annals of Botany (Oxford) 98 (1): 41–47. doi:10.1093/aob/mcl082. PMC 2803533. PMID 16687431. 
  60. ^ Gon g, L.; Pachner, M.; Kalai, K.; Lelley, T. (November 2008). "SSR-based genetic linkage map of Cucurbita moschata and its Synteny With Cucurbita pepo". Genome 51 (11): 878–887. doi:10.1139/G08-072. PMID 18956020. 
  61. ^ Gong, L.; Stift, G.; Koffler, R.; Pachner, M.; Lelley, T. (June 2008). "Microsatellites for the genus Cucurbita and an SSR-based genetic linkage map of Cucurbita pepo L.". Theoretical and Applied Genetics 117 (1): 37–48. doi:10.1007/s00122-008-0750-2. PMID 18379753. 
  62. ^ Linnaeus, Carl (1753). "Cucurbita". Species Plantarum 2. Stockholm: Laurentius Salvius. p. 2010. 
  63. ^ "Cucurbita". The Linnaean Plant Name Typification Project. Natural History Museum. Retrieved November 4, 2013. 
  64. ^ "Cross Pollination in Cucurbits" (PDF). Purdue Agriculture. Retrieved September 1, 2013. 
  65. ^ GRIN. "GRIN Species Records of genus Cucurbita". Taxonomy for Plants. National Germplasm Resources Laboratory, Beltsville, Maryland: USDA, ARS, National Genetic Resources Program. Retrieved September 1, 2013. 
  66. ^ "Cucurbita". Integrated Taxonomic Information System. Retrieved September 1, 2013. 
  67. ^ Andres, Thomas C. (1987). "Relationship of Cucurbita scabridifolia to C. foetidissima and C. pedatifolia: A Case of Natural Interspecific Hybridization". Cucurbit Genetics Cooperative Report (Raleigh, NC: North Carolina State University) 10: 74–75. 
  68. ^ Bailey, Liberty Hyde (1943). "Species of Cucurbita". Gentes Herbarum (Ithaca, NY) 6: 267–322. 
  69. ^ Bemis, W. P.; Rhodes, A. M.; Whitaker, Thomas W.; Carmer, S. G. (1970). "Numerical Taxonomy Applied to Cucurbita Relationships". American Journal of Botany (Botanical Society of America) 57 (4): 404–412. doi:10.2307/2440868. JSTOR 2440868.  edit
  70. ^ "Cucurbita maxima Origin/ Habitat". University of Wisconsin. 2007. Retrieved September 2, 2013. 
  71. ^ "Holotype of Cucurbita maxima Duchesne var. triloba Millán [family CUCURBITACEAE]". JSTOR. Retrieved October 3, 2013. 
  72. ^ López-Anido, F.; Cravero, V.; Asprelli, P.; Cointry, E.; Firpo, I.; García, S. M. (2003). "Inheritance of Immature Fruit Color in Cucurbita maxima var. Zapallito (Carrière) Millán" (PDF). Cucurbit Genetics Cooperative Report (Raleigh, NC: North Carolina State University) 26: 48–50. 
  73. ^ "Holotype of Cucurbita maxima Duchesne var. zipinka Millán [family CUCURBITACEAE]". JSTOR. Retrieved October 3, 2013. 
  74. ^ a b Robinson, Richard Warren; Decker-Walters, D. S. (1997). Cucurbits. Oxfordshire, UK: CAB International. pp. 71–83. ISBN 978-0-8519-9133-7. 
  75. ^ Wessel-Beaver, Linda (2000). "Evidence for the Center of Diversity of Cucurbita moschata in Colombia". Cucurbit Genetics Cooperative Report (Raleigh, NC: North Carolina State University) 23: 54–55. 
  76. ^ Kirkpatrick, Kurt J.; Wilson, Hugh D. (1988). "Interspecific Gene Flow in Cucurbita: C. texana vs. C. pepo". American Journal of Botany (Botanical Society of America) 75 (4): 519–527. JSTOR 2444217.  edit
  77. ^ a b c Andres, Thomas C. (1987). "Cucurbita fraterna, the Closest Wild Relative and Progenitor of C. pepo". Cucurbit Genetics Cooperative Report (Raleigh, NC: North Carolina State University) 10: 69–71. 
  78. ^ Soltis, Douglas E.; Soltis, Pamela S. Isozymes in Plant Biology. London: Dioscorodes Press. p. 176. ISBN 978-0-412-36500-3. 
  79. ^ a b c d e f g h i j Paris, Harry S. (1986). "A Proposed Subspecific Classification for Cucurbita pepo". Phytologia (Bronx Park) 61 (3): 133–138. 
  80. ^ a b c d e f g h i "Cucurbita pepo". Missouri Botanical Garden. Retrieved September 2, 2013. 
  81. ^ a b c d e f g h i Heistinger, Andrea (2013). The Manual of Seed Saving: Harvesting, Storing, and Sowing Techniques for Vegetables, Herbs, and Fruits. Portland, OR: Timber Press. p. 278. ISBN 978-1-60469-382-9. 
  82. ^ a b Decker, Deena S.; Wilson, Hugh D. (1987). "Allozyme Variation in the Cucurbita pepo Complex: C. pepo var. ovifera vs. C. texana". Systematic Botany (American Society of Plant Taxonomists) 12 (2): 263–273. doi:10.2307/2419320. JSTOR 2419320.  edit
  83. ^ a b Fürnkranz, Michael; Lukesch, Birgit; Müller, Henry; Huss, Herbert; Grube, Martin; Berg, Gabriele (2012). "Microbial Diversity Inside Pumpkins: Microhabitat-Specific Communities Display a High Antagonistic Potential Against Phytopathogens". Microbial Ecology (Springer) 63 (2): 418–428. doi:10.2307/41412429. JSTOR 41412429.  edit
  84. ^ Lim, Tong Kwee (2012). Edible Medicinal And Non-Medicinal Plants: Volume 2, Fruits. New York: Springer. p. 283. ISBN 978-94-007-1763-3. 
  85. ^ Lazos, E. S. (July 1992). "Certain Functional Properties of Defatted Pumpkin Seed Flour". Plant Food for Human Nutrition 42 (3): 257–273. doi:10.1007/bf02193934. PMID 1502127. 
  86. ^ a b Bean, Lowell J.; Saubel, Katherine Siva (1972). Temalpakh: Cahuilla Indian Knowledge and Usage of Plants. Morongo Indian Reservation, Riverside County, CA: Malki Museum Press. p. 57. ISBN 978-0-9390-4624-9. 
  87. ^ Richardson, R. W. "Squash and Pumpkin" (PDF). United States Department of Agriculture, Agricultural Research Service, National Plant Germplasm System. Retrieved September 2, 2013. 
  88. ^ "Spiced Pumpkin Butter". Better Homes and Gardens. Retrieved January 12, 2014. 
  89. ^ Lynch, Rene (October 1, 2013). "Pumpkin Bread and 18 Other Pumpkin Recipes You Must Make Now". Los Angeles Times. Retrieved October 8, 2013. 
  90. ^ Bemis, W. P. (1978). "The Versatility of the Feral Buffalo Gourd, Cucurbita foetidissima HBK". Cucurbit Genetics Cooperative Report (Raleigh, NC: North Carolina State University) 1: 25. 
  91. ^ Mihranian, Valentine H.; Abou-Chaar, Charles I. (1968). "Extraction, Detection, and Estimation of Cucurbitin in Cucurbita seeds". Lloydia (American Society of Pharmacognosy) 31 (1): 23–29. 
  92. ^ Tang, Weici; Eisenbrand, Gerhard (1992). "Cucurbita moschata Duch.". Chinese Drugs of Plant Origin. Berlin: Springer-Verlag. pp. 399–400. doi:10.1007/978-3-642-73739-8_51. ISBN 978-3-642-73741-1. 
  93. ^ Xiao, S. H.; Keiser, J.; Chen, M. G.; Tanner, M.; Utzinger, J. (2010). "Research and Development of antischistosomal drugs in the People's Republic of China a 60-year review". Advances in Parasitology 73: 231–295. doi:10.1016/S0065-308X(10)73009-8. PMID 20627145. 
  94. ^ "Full Report (All Nutrients): 11422, Pumpkin, raw". Agricultural Research Service, United States Department of Agriculture. Retrieved September 21, 2013. 
  95. ^ a b "Pumpkin". 2009. Retrieved September 1, 2013. 
  96. ^ Pierce, Andrea (1999). The Apha Practical Guide to Natural Medicines. New York: Stonesong Press, William Morrow & Company. pp. 212–214. ISBN 0-688-16151-0. 
  97. ^ Yadav, M.; Jain, S.; Tomar, R.; Prasad, G. B.; Yadav, H. (December 2010). "Medicinal and Biological Potential of Pumpkin: An Updated Review". Nutritional Research Reviews 23 (2): 184–190. doi:10.1017/S0954422410000107. PMID 21110905. 
  98. ^ Mansour, Esam H.; Dworschák, Erno; Lugasi, Andrea; Barna, Barna; Gergely, Anna (1993). "Nutritive Value of Pumpkin (Cucurbita Pepo Kakai 35) Seed Products". Journal of the Science of Food and Agriculture 61 (1): 73–78. doi:10.1002/jsfa.2740610112. 
  99. ^ Kreider, Richard B.; Leutholtz, Brian C.; Katch, Frank I.; Katch, Victor L. (2009). Exercise and Sport Nutrition. Santa Barbara, CA: Fitness Technologies Press. p. 285. ISBN 978-0-974-29656-2. 
  100. ^ "Pumpkin Nutrition". University of Illinois Extenstion. Retrieved October 20, 2013. 
  101. ^ "What's So Great About Winter Squash?" (PDF). University of the District of Columbia. Retrieved October 22, 2013. 
  102. ^ a b Preedy, Victor R.; Watson, Ronald Ross; Patel, Vinwood B. (2011). Nuts and Seeds in Health and Disease Prevention. London: Academic Press. p. 936. ISBN 978-0-12-375688-6. 
  103. ^ Barbieri, L.; Polito, L.; Bolognesi, A.; Ciani, M.; Pelosi, E.; Farini, V.; Jha, A. K.; Sharma, N.; Vivanco, J. M.; Chambery, A.; Parente, A.; Stirpe, F. (May 2006). "Ribosome-inactivating Proteins in Edible Plants and Purification and Characterization of a New Ribosome-inactivating Protein From Cucurbita moschata". Biochimica et Biophysica Acta 1760 (5): 783–792. doi:10.1016/j.bbagen.2006.01.002. PMID 16564632. 
  104. ^ Hou, Xiaomin; Meehan, Edward J.; Xie, Jieming; Huang, Mingdong; Chen, Minghuang; Chen, Liqing (October 2008). "Atomic resolution structure of cucurmosin, a novel type 1 ribosome-inactivating protein from the sarcocarp of Cucurbita moschata". Journal of Structural Biology 164 (1): 81–87. doi:10.1016/j.jsb.2008.06.011. 
  105. ^ a b Tallamy, Douglas W.; Krischik, Vera A. (1989). "Variation and Function of Cucurbitacins in Cucurbita: An Examination of Current Hypotheses". The American Naturalist (The University of Chicago Press) 133 (6): 766–786. doi:10.1086/284952. JSTOR 2462036.  edit
  106. ^ Chen, Jian Chao; Chiu, Ming Hua; Nie, Rui Lin; Cordell, Geoffrey A.; Qui, Samuel X. (2005). "Cucurbitacins and Cucurbitane Glycosides: Structures and Biological Activities". Natural Product Reports 22 (5): 386–399. doi:10.1039/B418841C. PMID 16010347. 
  107. ^ McAuslane, Heather J.; Elmstrom, Gary W. (June 1996). "Resistance in Germplasm of Cucurbita pepo to Silverleaf, a Disorder Associated with Bemisia argentifolii (Homoptera: Aleyrodidae)". The Florida Entomologist (Lutz, FL: Florida Entomological Society) 79 (2): 206–221. doi:10.2307/3495818. JSTOR 3495818.  edit
  108. ^ a b "Vegetable Pumpkin". University of Illinois Extension. Retrieved October 20, 2013. 
  109. ^ "Common name: Squash Bug". University of Florida. Retrieved October 26, 2013. 
  110. ^ a b "Mosaic Diseases of Cucurbits" (PDF). University of Illinois. Retrieved October 26, 2013. 
  111. ^ a b "Virus Diseases of Cucurbit Crops" (PDF). Department of Agriculture, Government of Western Australia. Retrieved October 26, 2013. 
  112. ^ Roossinck, Marilyn J.; Palukaitis, Peter (1990). "Rapid Induction and Severity of Symptoms in Zucchini Squash (Cucurbita pepo) Map to RNA 1 of Cucumber Mosaic Virus" (PDF). Molecular Plant-Microbe Interactions 3 (3): 188–192. doi:10.1094/mpmi-3-188. 
  113. ^ Havelda, Zoltan; Maule, Andrew J. (October 2000). "Complex Spatial Responses to Cucumber Mosaic Virus Infection in Susceptible Cucurbita pepo Cotyledons". Plant Cell 12 (10): 1975–1986. doi:10.1105/tpc.12.10.1975. PMC 149134. PMID 11041891. 
  114. ^ "Virus Diseases of Cucurbits". Cornell University. October 1984. Retrieved October 26, 2013. 
  115. ^ Provvidenti, R.; Gonsalves, D. (May 1984). "Occurrence of Zucchini Yellow Mosaic Virus in Cucurbits from Connecticut, New York, Florida, and California" (PDF). Plant Disease 68 (5). pp. 443–446. doi:10.1094/pd-69-443. ISSN 0191-2917. 
  116. ^ "Squash". Texas A&M University. Retrieved October 26, 2013. 
  117. ^ Salama, El-Sayed A.; Sill, Jr., W. H. (1968). "Resistance to Kansas Squash Mosaic Virus Strains Among Cucurbita Species". Transactions of the Kansas Academy of Science 71 (1): 62–68. JSTOR 3627399.  edit
  118. ^ a b Geisler, Malinda (May 2012). "Squash". Agricultural Marketing Resource Center, Iowa State University. Retrieved October 13, 2013. 
  119. ^ a b c d e "Pumpkins, Squash, and Gourds". Food and Agriculture Organization of the United Nations. 2012. Retrieved October 13, 2013. 

External links[edit]