Cucurbita maxima Zapallo Plomo semillería Costanzi - vine detail decumbent shoot and tendril.j
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''Cucurbita'' (
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
for
gourd 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. One of the ear ...
) is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of herbaceous
fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...
s in the gourd
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
,
Cucurbitaceae The Cucurbitaceae, also called cucurbits or the gourd family, are a plant family consisting of about 965 species in around 95 genera, of which the most important to humans are: *''Cucurbita'' – squash, pumpkin, zucchini, some gourds *'' Lagen ...
(also known as ''cucurbits'' or ''cucurbi''), native to the
Andes The Andes, Andes Mountains or Andean Mountains (; ) are the longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range is long, wide (widest between 18°S – 20°S ...
and
Mesoamerica Mesoamerica is a historical region and cultural area in southern North America and most of Central America. It extends from approximately central Mexico through Belize, Guatemala, El Salvador, Honduras, Nicaragua, and northern Costa Rica ...
. 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 Variety may refer to: Arts and entertainment Entertainment formats * Variety (radio) * Variety show, in theater and television Films * ''Variety'' (1925 film), a German silent film directed by Ewald Andre Dupont * ''Variety'' (1935 film), ...
, and local parlance. Other kinds of gourd, also called bottle-gourds, are native to Africa and belong to the genus ''
Lagenaria ''Lagenaria'' is a genus of gourd-bearing vines in the squash family (Cucurbitaceae). ''Lagenaria'' contains six species, all of which are indigenous to tropical Africa.tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to confli ...
. These other gourds are used as utensils or vessels, and their young fruits are eaten much like those of the ''Cucurbita'' species. Most ''Cucurbita'' species are herbaceous vines that grow several meters in length and have
tendril In botany, a tendril is a specialized stem, leaf or petiole with a threadlike shape used by climbing plants for support and attachment, as well as cellular invasion by parasitic plants such as '' Cuscuta''. There are many plants that have tend ...
s, 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 bee A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the current cosm ...
s, also visit. There is debate about the
taxonomy Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
of the genus, as the number of accepted species varies from 13 to 30. The five
domesticated Domestication is a sustained multi-generational relationship in which humans assume a significant degree of control over the reproduction and care of another group of organisms to secure a more predictable supply of resources from that group. A ...
species are ''
Cucurbita argyrosperma ''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, ...
'', '' C. ficifolia'', '' C. maxima'', '' C. moschata'', and '' C. pepo''. All of these can be treated as
winter squash 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 ...
because the full-grown fruits can be stored for months; however, ''C. pepo'' includes some
cultivar A cultivar is a type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture ...
s that are better used only as
summer squash Summer squash are squashes that are harvested when immature, while the rind is still tender and edible. Nearly all summer squashes are varieties of ''Cucurbita pepo'', although not all ''Cucurbita pepo'' are considered summer squashes. Most summ ...
. The fruits of the genus ''Cucurbita'' are good sources of nutrients, such as
vitamin A Vitamin A is a fat-soluble vitamin and an essential nutrient for humans. It is a group of organic compounds that includes retinol, retinal (also known as retinaldehyde), retinoic acid, and several provitamin A carotenoids (most notably ...
and
vitamin C Vitamin C (also known as ascorbic acid and ascorbate) is a water-soluble vitamin found in citrus and other fruits and vegetables, also sold as a dietary supplement and as a topical 'serum' ingredient to treat melasma (dark pigment spots) ...
, among other nutrients according to species. The fruits have many culinary uses including
pumpkin pie Pumpkin pie is a dessert pie with a spiced, pumpkin-based custard filling. The pumpkin and pumpkin pie are both a symbol of harvest time, and pumpkin pie is generally eaten during the fall and early winter. In the United States and Canada it is u ...
, biscuits, bread, desserts, puddings, beverages, and soups. Although botanical fruits, ''Cucurbita'' gourds such as squash are typically cooked and eaten as
vegetable Vegetables are parts of plants that are consumed by humans or other animals as food. The original meaning is still commonly used and is applied to plants collectively to refer to all edible plant matter, including the flowers, fruits, stems, ...
s. Pumpkins see more varied use, and are eaten both as vegetables and as desserts such as
pumpkin pie Pumpkin pie is a dessert pie with a spiced, pumpkin-based custard filling. The pumpkin and pumpkin pie are both a symbol of harvest time, and pumpkin pie is generally eaten during the fall and early winter. In the United States and Canada it is u ...
.


Description

''Cucurbita'' species fall into two main groups. The first group are
annual Annual may refer to: *Annual publication, periodical publications appearing regularly once per year ** Yearbook ** Literary annual *Annual plant *Annual report *Annual giving *Annual, Morocco, a settlement in northeastern Morocco *Annuals (band), ...
or short-lived
perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wid ...
vines and are
mesophytic Mesophytes are terrestrial plants which are neither adapted to particularly dry nor particularly wet environments. An example of a mesophytic habitat would be a rural temperate meadow, which might contain goldenrod, clover, oxeye daisy, and ''Rosa m ...
, i.e. they require a more or less continuous water supply. The second group are
perennials A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also widel ...
growing in arid zones and so are
xerophytic A xerophyte (from Greek ξηρός ''xeros'' 'dry' + φυτόν ''phuton'' 'plant') is a species of plant that has adaptations to survive in an environment with little liquid water, such as a desert such as the Sahara or places in the Alps or t ...
, tolerating dry conditions. Cultivated ''Cucurbita'' species were derived from the first group. Growing in height or length, the plant stem produces
tendril In botany, a tendril is a specialized stem, leaf or petiole with a threadlike shape used by climbing plants for support and attachment, as well as cellular invasion by parasitic plants such as '' Cuscuta''. There are many plants that have tend ...
s 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. The morphological variation in the species ''C. pepo'' and ''C. maxima'' is so vast that its various subspecies and cultivars have been misidentified as totally separate species. The typical cultivated ''Cucurbita'' species has five-lobed or palmately divided leaves with long petioles, 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
trichome Trichomes (); ) are fine outgrowths or appendages on plants, algae, lichens, and certain protists. They are of diverse structure and function. Examples are hairs, glandular hairs, scales, and papillae. A covering of any kind of hair on a p ...
s, which are often hardened and sharp. Spring-like tendrils grow from each
node In general, a node is a localized swelling (a "knot") or a point of intersection (a vertex). Node may refer to: In mathematics * Vertex (graph theory), a vertex in a mathematical graph *Vertex (geometry), a point where two or more curves, lines ...
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 Puberty is the process of physical changes through which a child's body matures into an adult body capable of sexual reproduction. It is initiated by hormonal signals from the brain to the gonads: the ovaries in a girl, the testes in a boy. I ...
. ''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. The species are monoecious, with unisexual male (
staminate The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filame ...
) 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 together so that there appears to be one. Female flowers have thick
pedicels In botany, a pedicel is a stem that attaches a single flower to the inflorescence. Such inflorescences are described as ''pedicellate''. Description Pedicel refers to a structure connecting a single flower to its inflorescence. In the absenc ...
, and an
inferior ovary In the flowering plants, an ovary is a part of the female reproductive organ of the flower or gynoecium. Specifically, it is the part of the pistil which holds the ovule(s) and is located above or below or at the point of connection with the ba ...
with 3–5 stigmas that each have two lobes. The female flowers of ''C. argyrosperma'' and ''C. ficifolia'' have larger corollas than the male flowers. Female flowers of ''C. pepo'' have a small calyx, but the calyx of ''C. moschata'' male flowers is comparatively short. ''Cucurbita'' fruits are large and fleshy. 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 In angiosperms, a hypanthium or floral cup is a structure where basal portions of the calyx, the corolla, and the stamens form a cup-shaped tube. It is sometimes called a floral tube, a term that is also used for corolla tube and calyx tube. It ...
tissue forming an
exocarp Fruit anatomy is the plant anatomy of the internal structure of fruit. Fruits are the mature ovary or ovaries of one or more flowers. They are found in three main anatomical categories: aggregate fruits, multiple fruits, and simple fruits. Aggr ...
around the ovary, and a fleshy interior composed of
mesocarp Fruit anatomy is the plant anatomy of the internal structure of fruit. Fruits are the mature ovary or ovaries of one or more flowers. They are found in three main anatomical categories: aggregate fruits, multiple fruits, and simple fruits. Agg ...
and
endocarp Fruit anatomy is the plant anatomy of the internal structure of fruit. Fruits are the mature ovary or ovaries of one or more flowers. They are found in three main anatomical categories: aggregate fruits, multiple fruits, and simple fruits. Aggr ...
. The term "pepo" is used primarily for Cucurbitaceae fruits, where this fruit type is common, but the fruits of '' Passiflora'' and ''
Carica ''Carica'' is a genus of flowering plants in the family Caricaceae including the papaya (''C. papaya'' syn. ''C. peltata'', ''C. posoposa''), a widely cultivated fruit tree native to the American tropics. The genus was formerly treated as incl ...
'' are sometimes also pepos. 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
cotyledon A cotyledon (; ; ; , gen. (), ) is a significant part of the embryo within the seed of a plant, and is defined as "the embryonic leaf in seed-bearing plants, one or more of which are the first to appear from a germinating seed." The num ...
s. Fruit size varies considerably: wild fruit specimens can be as small as and some domesticated specimens can weigh well over . The current world record was set in 2014 by Beni Meier of Switzerland with a pumpkin.


Taxonomy

''Cucurbita'' was formally described in a way that meets the requirements of modern botanical nomenclature by
Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
in his ''
Genera Plantarum ''Genera Plantarum'' is a publication of Swedish naturalist Carl Linnaeus (1707–1778). The first edition was issued in Leiden, 1737. The fifth edition served as a complementary volume to ''Species Plantarum'' (1753). Article 13 of the Internati ...
'', the fifth edition of 1754 in conjunction with the 1753 first edition of '' Species Plantarum''. ''Cucurbita pepo'' is the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specime ...
of the genus. 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 Watermelon (''Citrullus lanatus'') is a flowering plant species of the Cucurbitaceae family and the name of its edible fruit. A scrambling and trailing vine-like plant, it is a highly cultivated fruit worldwide, with more than 1,000 varieti ...
'') and ''C. lagenaria'' (now ''
Lagenaria siceraria Calabash (; ''Lagenaria siceraria''), also known as bottle gourd, white-flowered gourd, long melon, birdhouse gourd, New Guinea bean, Tasmania bean, and opo squash, is a vine grown for its fruit. It can be either harvested young to be consumed ...
'') (both are not ''Cucurbita'' but are in the family Cucurbitaceae. The ''Cucurbita digitata'', ''C. foetidissima'', ''C. galeotti'', and ''C. pedatifolia''
species group In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
s are
xerophytes A xerophyte (from Greek ξηρός ''xeros'' 'dry' + φυτόν ''phuton'' 'plant') is a species of plant that has adaptations to survive in an environment with little liquid water, such as a desert such as the Sahara or places in the Alps or the ...
, 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. The five domesticated species are mostly isolated from each other by sterility barriers and have different physiological characteristics. 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. The buffalo gourd (''C. foetidissima'') has been used as an intermediary, as it can be crossed with all the common ''Cucurbita''. Various taxonomic treatments have been proposed for ''Cucurbita'', ranging from 13 to 30 species. 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: * '' C. argyrosperma'' ( synonym ''C. mixta'') – cushaw pumpkin; origin: Mexico ** '' C. kellyana'', origin: Pacific coast of western Mexico ** '' C. palmeri'', origin: Pacific coast of northwestern Mexico ** '' C. sororia'', origin: Pacific coast Mexico to Nicaragua, northeastern Mexico * '' C. digitata'' – fingerleaf gourd; origin: southwestern United States (USA), northwestern Mexico ** '' C. californica'' ** '' C. cordata'' ** '' C. cylindrata'' ** '' C. palmata'' * '' C. ecuadorensis'', origin: Ecuador's Pacific coast * '' C. ficifolia'' – figleaf gourd, chilacayote, alcayota; origin: Mexico, Panama, northern Chile and Argentina * '' C. foetidissima'' – stinking gourd, buffalo gourd; origin: Mexico ** '' C. scabridifolia'', likely a
natural hybrid In biology, a hybrid is the offspring resulting from combining the qualities of two organisms of different breeds, varieties, species or genera through sexual reproduction. Hybrids are not always intermediates between their parents (such as in ...
of ''C. foetidissima'' and ''C. pedatifolia'' * '' C. galeottii'', little known; origin:
Oaxaca Oaxaca ( , also , , from nci, Huāxyacac ), officially the Free and Sovereign State of Oaxaca ( es, Estado Libre y Soberano de Oaxaca), is one of the 32 states that compose the political divisions of Mexico, Federative Entities of Mexico. It is ...
, Mexico * '' C. lundelliana'', origin: Mexico, Guatemala, Belize * '' C. maxima'' – winter squash, pumpkin; origin: Argentina, Bolivia, Ecuador ** '' C. andreana'', origin – Argentina * '' C. moschata'' – butternut squash, 'Dickinson' pumpkin, golden cushaw; origin: Bolivia, Colombia, Ecuador, Mexico, Panama, Puerto Rico, Venezuela * '' C. okeechobeensis'', origin: Florida ** '' C. martinezii'', origin: Mexican Gulf Coast and foothills * '' C. pedatifolia'', origin: Querétaro, Mexico ** '' C. moorei'' * '' C. pepo'' – field pumpkin, summer squash, zucchini, vegetable marrow, courgette, acorn squash; origin: Mexico, USA ** '' C. fraterna'', origin:
Tamaulipas Tamaulipas (), officially the Free and Sovereign State of Tamaulipas ( es, Estado Libre y Soberano de Tamaulipas), is a state in the northeast region of Mexico; one of the 31 states which, along with Mexico City, comprise the 32 Federal Entiti ...
and Nuevo León, Mexico ** '' C. texana'', origin:
Texas Texas (, ; Spanish: ''Texas'', ''Tejas'') is a state in the South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2020, it is the second-largest U.S. state by ...
, USA * '' C. radicans'' – calabacilla, calabaza de coyote; origin: Central Mexico ** '' C. gracilior'' The taxonomy by Nee closely matches the species groupings reported in a pair of studies by a botanical team led by Rhodes and Bemis in 1968 and 1970 based on statistical groupings of several
phenotypic In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology or physical form and structure, its developmental processes, its biochemical and physiological proper ...
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: * ''C. digitata'', ''C. palmata'', ''C. californica'', ''C. cylindrata'', ''C. cordata'' * ''C. martinezii'', ''C. okeechobeensis'', ''C. lundelliana'' * ''C. sororia'', ''C. gracilior'', ''C. palmeri''; ''C. argyrosperma'' (reported as ''C. mixta'') was considered close to the three previous species * ''C. maxima'', ''C. andreana'' * ''C. pepo'', ''C. texana'' * ''C. moschata'', ''C. ficifolia'', ''C. pedatifolia'', ''C. foetidissima'', and ''C. ecuadorensis'' were placed in their own separate species groups as they were not considered significantly close to any of the other species studied.


Phylogeny

The full
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spe ...
of this genus is unknown, and research was ongoing in 2014. The following
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
of ''Cucurbita'' phylogeny is based upon a 2002 study of mitochondrial DNA by Sanjur and colleagues.


Reproductive biology

All species of ''Cucurbita'' have 20 pairs of
chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
s. Many North and Central American species are visited by specialist pollinators in the apid tribe Eucerini, especially the genera ''
Peponapis ''Peponapis'' is a genus of bees belonging to the family Apidae. The species of this genus are found in America. Species: *'' Peponapis apiculata'' *'' Peponapis atrata'' *''Peponapis azteca'' *''Peponapis citrullina'' *'' Peponapis crassi ...
'' and '' Xenoglossa'', and these
squash bee 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 (pollen specialists) on the plant genus ''Cucurbita'' and closely relate ...
s can be crucial to the flowers producing fruit after pollination. 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, an effect called xenia. Competitively grown specimens are therefore often hand-pollinated to maximize the number of seeds in the fruit, which increases the fruit size; this pollination requires skilled technique. Seedlessness is known to occur in certain cultivars of ''C. pepo''. The most critical factors in flowering and fruit set are physiological, having to do with the age of the plant and whether it already has developing fruit. The plant hormones ethylene and auxin are key in fruit set and development. 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", and male flowers are more frequent, an effect that appears due to reduced natural ethylene production within the plant stem.
Ethephon Ethephon is a plant growth regulator. Mechanism of action Upon metabolism by the plant, it is converted into ethylene, a potent regulator of plant growth and ripeness. It is also a butyrylcholinesterase inhibitor. Uses in various crops Etheph ...
, a plant growth regulator product that is converted to ethylene after metabolism by the plant, can be used to increase fruit and seed production. The plant hormone
gibberellin Gibberellins (GAs) are plant hormones that regulate various developmental processes, including stem elongation, germination, dormancy, flowering, flower development, and leaf and fruit senescence. GAs are one of the longest-known classes of plan ...
, 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. Gibberellin is also involved in other developmental processes of plants such as seed and stem growth.


Germination and seedling growth

Seeds with maximum germination potential develop (in ''C. moschata'') by 45 days after
anthesis Anthesis is the period during which a flower is fully open and functional. It may also refer to the onset of that period. The onset of anthesis is spectacular in some species. In ''Banksia'' species, for example, anthesis involves the extension ...
, and seed weight reaches its maximum 70 days after anthesis. Some varieties of ''C. pepo'' germinate best with eight hours of sunlight daily and a planting depth of . Seeds planted deeper than are not likely to germinate. 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. 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. 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 Phytic acid is a six-fold dihydrogenphosphate ester of inositol (specifically, of the ''myo'' isomer), also called inositol hexakisphosphate (IP6) or inositol polyphosphate. At physiological pH, the phosphates are partially ionized, resulting in ...
forms in seed tissues as spherical crystalline intrusions in protein bodies called globoids. Along with other nutrients, phytate is used completely during seedling growth. Heavy metal contamination, including
cadmium Cadmium is a chemical element with the symbol Cd and atomic number 48. This soft, silvery-white metal is chemically similar to the two other stable metals in group 12, zinc and mercury. Like zinc, it demonstrates oxidation state +2 in most of ...
, has a significant negative impact on plant growth. ''Cucurbita'' plants grown in the spring tend to grow larger than those grown in the autumn.


Distribution and habitat

Archaeological investigations have found evidence of domestication of Cucurbita going back over 8,000 years from the very southern 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. Of the 27 species that Nee delineates, five are domesticated. Four of them, ''C. argyrosperma'', ''C. ficifolia'', ''C. moschata'', and ''C. pepo'', originated and were domesticated in
Mesoamerica Mesoamerica is a historical region and cultural area in southern North America and most of Central America. It extends from approximately central Mexico through Belize, Guatemala, El Salvador, Honduras, Nicaragua, and northern Costa Rica ...
; for the fifth, ''C. maxima'', these events occurred in South America. 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 bigger but fewer in number. 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 is a cross between the scallop and the pumpkin. ''C. argyrosperma'' is not as widespread as the other species. The wild form ''C. a.'' subsp. ''sororia'' is found from
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
to
Nicaragua Nicaragua (; ), officially the Republic of Nicaragua (), is the largest country in Central America, bordered by Honduras to the north, the Caribbean to the east, Costa Rica to the south, and the Pacific Ocean to the west. Managua is the countr ...
, and cultivated forms are used in a somewhat wider area stretching from Panama to the southeastern United States. It was probably bred for its seeds, which are large and high in
oil An oil is any nonpolar chemical substance that is composed primarily of hydrocarbons and is hydrophobic (does not mix with water) & lipophilic (mixes with other oils). Oils are usually flammable and surface active. Most oils are unsaturated ...
and
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
, 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 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. ''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 in areas with heavy rainfall. It does not hybridize well with the other cultivated species as it has significantly different enzymes and chromosomes. ''C. maxima'' originated in South America over 4,000 years ago, 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. ''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. Types of ''C. maxima'' include ''triloba'', ''zapallito'', ''zipinka'', Banana, Delicious, Hubbard, Marrow (''C. maxima'' Marrow), Show, and Turban. ''C. moschata'' is native to Latin America, but the precise location of origin is uncertain. 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 wide. It generally grows at low elevations in hot climates with heavy rainfall, but some varieties have been found above . Groups of ''C. moschata'' include Cheese, Crookneck (''C. moschata''), and Bell. ''C. pepo'' is one of the oldest, if not the oldest, domesticated species with the oldest known locations being
Oaxaca Oaxaca ( , also , , from nci, Huāxyacac ), officially the Free and Sovereign State of Oaxaca ( es, Estado Libre y Soberano de Oaxaca), is one of the 32 states that compose the political divisions of Mexico, Federative Entities of Mexico. It is ...
, Mexico, 8,000–10,000 years ago, and
Ocampo, Tamaulipas Ciudad Ocampo in Ocampo Municipality in the Mexican state of Tamaulipas was founded on May 19, 1749 as Villa of Santa Bárbara; the town became known as Ocampo in 1869. The credit for the founding of Santa Barbara is given to Don José Escandón y ...
, Mexico, about 7,000 years ago. It is known to have appeared in
Missouri Missouri is a state in the Midwestern region of the United States. Ranking 21st in land area, it is bordered by eight states (tied for the most with Tennessee): Iowa to the north, Illinois, Kentucky and Tennessee to the east, Arkansas t ...
, United States, at least 4,000 years ago. Debates about the origin of ''C. pepo'' have been on-going since at least 1857. 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 A feral () animal or plant is one that lives in the wild but is descended from domesticated individuals. As with an introduced species, the introduction of feral animals or plants to non-native regions may disrupt ecosystems and has, in some ...
''C. pepo''. A more recent theory by botanist Thomas Andres in 1987 is that descendants of ''C. fraterna'' hybridized with ''C. texana'', 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. ''C. pepo'' may have appeared in the Old World before moving from Mexico into South America. It is found from sea level to slightly above . Leaves have 3–5 lobes and are wide. All the subspecies, varieties, and
cultivar A cultivar is a type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture ...
s are interfertile. In 1986 Paris proposed a revised taxonomy of the edible cultivated ''C. pepo'' based primarily on the shape of the fruit, with eight groups . All but a few ''C. pepo'' cultivars can be included in these groups. There is one non-edible cultivated variety: ''C. pepo'' var. ''ovifera''.


History and domestication

The ancestral species of the genus ''Cucurbita'' were present in the Americas before the arrival of humans, and are native to the
Americas The Americas, which are sometimes collectively called America, are a landmass comprising the totality of North and South America. The Americas make up most of the land in Earth's Western Hemisphere and comprise the New World. Along with th ...
. The likely center of origin is southern Mexico, spreading south through what is now known as Mesoamerica, on into South America, and north to what is now the southwestern United States. Evolutionarily speaking, the genus is relatively recent in origin, dating back only to the
Holocene The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togeth ...
, whereas the family Cucurbitaceae, in the shape of seeds similar to ''
Bryonia ''Bryonia'' is a genus of flowering plants in the gourd family. Bryony is its best-known common name. They are native to western Eurasia and adjacent regions, such as North Africa, the Canary Islands and South Asia. Description and ecology B ...
'', dates to the
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pal ...
. Recent genomic studies support the idea that the ''Cucurbita'' genus underwent a whole-genome duplication event, increasing their number of chromosomes and accelerating the rate at which their genomes evolve compared to other cucurbits. No species within the genus is entirely genetically isolated. ''C. moschata'' can intercross with all the others, though the hybrid offspring may not themselves be fertile unless they become
polyploid Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei ( eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contain ...
. The genus was part of the culture of almost every native peoples group from southern South America to southern Canada. Modern-day cultivated ''Cucurbita'' are not found in the wild. Genetic studies of the
mitochondrial gene Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial D ...
'' nad1'' show there were at least six independent domestication events of ''Cucurbita'' separating domestic species from their wild ancestors. Species native to North America include '' C. digitata'' (calabazilla), and ''C. foetidissima'' (buffalo gourd), '' C. palmata'' (coyote melon), and ''C. pepo''. 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 ''Lagenaria'' is a genus of gourd-bearing vines in the squash family (Cucurbitaceae). ''Lagenaria'' contains six species, all of which are indigenous to tropical Africa.tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to confli ...
. 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 Maize ( ; ''Zea mays'' subsp. ''mays'', from es, maíz after tnq, mahiz), also known as corn (North American and Australian English), is a cereal grain first domesticated by indigenous peoples in southern Mexico about 10,000 years ago. The ...
and beans in the region by about 4,000 years. This evidence was found in the
Guilá Naquitz cave Guilá Naquitz Cave in Oaxaca, Mexico, is the site of early domestication of several food crops, including teosinte (an ancestor of maize), squash from the genus ''Cucurbita'', bottle gourds (''Lagenaria siceraria''), and beans. This site is the ...
in Oaxaca, Mexico, during a series of excavations in the 1960s and 1970s, possibly beginning in 1959. 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 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. During the same time frame, average rind thickness increased from . Recent genomic studies suggest that ''
Cucurbita argyrosperma ''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, ...
'' was domesticated in Mexico, in the region that is currently known as the state of Jalisco. Squash was domesticated first, followed by maize and then beans, becoming part of the Three Sisters agricultural system of
companion planting Companion planting in gardening and agriculture is the planting of different crops in proximity for any of a number of different reasons, including pest control, pollination, providing habitat for beneficial insects, maximizing use of space, a ...
. The
English English usually refers to: * English language * English people English may also refer to: Peoples, culture, and language * ''English'', an adjective for something of, from, or related to England ** English national ide ...
word "squash" derives from ''askutasquash'' (a green thing eaten raw), a word from the
Narragansett language Narragansett is an Algonquian language formerly spoken in most of what is today Rhode Island by the Narragansett people. It was closely related to the other Algonquian languages of southern New England like Massachusett and Mohegan-Pequot. T ...
, which was documented by Roger Williams, the founder of
Rhode Island Rhode Island (, like ''road'') is a state in the New England region of the Northeastern United States. It is the smallest U.S. state by area and the seventh-least populous, with slightly fewer than 1.1 million residents as of 2020, but it ...
, in his 1643 publication '' A Key Into the Language of America''. Similar words for squash exist in related languages of the Algonquian
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
.


Production

The family Cucurbitaceae has many species used as human food. ''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, 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. The seeds and fruits of most varieties can be stored for long periods of time, particularly the sweet-tasting winter varieties with their thick, inedible skins. Summer squash have a thin, edible skin. The seeds of both types can be roasted, eaten raw, made into
pumpkin seed oil Pumpkin seed oil is a culinary oil, used especially in central Europe. Culinary uses This oil is a culinary specialty from what used to be part of the Austria-Hungary, Austro-Hungarian Empire and is now southeastern Austria (Styria), eastern S ...
, ground into a flour or meal, or otherwise prepared. Squashes are primarily grown for the fresh food market. The
Food and Agriculture Organization of the United Nations The Food and Agriculture Organization of the United Nations (FAO)french: link=no, Organisation des Nations unies pour l'alimentation et l'agriculture; it, Organizzazione delle Nazioni Unite per l'Alimentazione e l'Agricoltura is an intern ...
(FAO) reported that the ranking of the top five squash-producing countries was stable between 2005 and 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 only additional 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. 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 percent of that from Mexico. Within the United States, the states producing the largest amounts are Florida, New York, California, and North Carolina.


Nutrients

As an example of ''Cucurbita'', raw summer squash is 94% water, 3% carbohydrates, and 1%
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
, with negligible
fat In nutrition, biology, and chemistry, fat usually means any ester of fatty acids, or a mixture of such compounds, most commonly those that occur in living beings or in food. The term often refers specifically to triglycerides (triple est ...
content (table). In a 100-gram reference serving, raw squash supplies of
food energy Food energy is chemical energy that animals (including humans) derive from their food to sustain their metabolism, including their muscular activity. Most animals derive most of their energy from aerobic respiration, namely combining the carbohy ...
and is rich in
vitamin C Vitamin C (also known as ascorbic acid and ascorbate) is a water-soluble vitamin found in citrus and other fruits and vegetables, also sold as a dietary supplement and as a topical 'serum' ingredient to treat melasma (dark pigment spots) ...
(20% of the
Daily Value The Reference Daily Intake (RDI) used in nutrition labeling on food and dietary supplement products in the U.S. and Canada is the daily intake level of a nutrient that is considered to be sufficient to meet the requirements of 97–98% of healthy ...
, DV), moderate in
vitamin B6 Vitamin B6 is one of the B vitamins, and thus an essential nutrient. The term refers to a group of six chemically similar compounds, i.e., " vitamers", which can be interconverted in biological systems. Its active form, pyridoxal 5′-phosp ...
and riboflavin (12–17% DV), but otherwise devoid of appreciable nutrient content (table), although the nutrient content of different ''Curcubita'' species may vary somewhat.
Pumpkin seed A pumpkin seed, also known in North America as a pepita (from the Mexican es, pepita de calabaza, "little seed of squash"), is the edible seed of a pumpkin or certain other cultivars of squash. The seeds are typically flat and asymmetrically ...
s contain
vitamin E Vitamin E is a group of eight fat soluble compounds that include four tocopherols and four tocotrienols. Vitamin E deficiency, which is rare and usually due to an underlying problem with digesting dietary fat rather than from a diet low in vi ...
,
crude protein Proteins are essential nutrients for the human body. They are one of the building blocks of body tissue and can also serve as a fuel source. As a fuel, proteins provide as much energy density as carbohydrates: 4 kcal (17 kJ) per gram; in cont ...
,
B vitamins B vitamins are a class of water-soluble vitamins that play important roles in cell metabolism and synthesis of red blood cells. Though these vitamins share similar names (B1, B2, B3, etc.), they are chemically distinct compounds that often coexi ...
and several
dietary minerals In the context of nutrition, a mineral is a chemical element required as an essential nutrient by organisms to perform functions necessary for life. However, the four major structural elements in the human body by weight (oxygen, hydrogen, carbon, ...
(see nutrition table at
pepita A pumpkin seed, also known in North America as a pepita (from the Mexican es, pepita de calabaza, "little seed of squash"), is the edible seed of a pumpkin or certain other cultivars of Squash (plant), squash. The seeds are typically flat and ...
). Also present in pumpkin seeds are unsaturated and
saturated Saturation, saturated, unsaturation or unsaturated may refer to: Chemistry * Saturation, a property of organic compounds referring to carbon-carbon bonds ** Saturated and unsaturated compounds **Degree of unsaturation ** Saturated fat or fatty ac ...
oils, palmitic, oleic and linoleic
fatty acids In chemistry, particularly in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated and unsaturated compounds#Organic chemistry, saturated or unsaturated. Most naturally occurring fatty acids have an B ...
, as well as
carotenoids Carotenoids (), also called tetraterpenoids, are yellow, orange, and red organic pigments that are produced by plants and algae, as well as several bacteria, and fungi. Carotenoids give the characteristic color to pumpkins, carrots, parsnips, co ...
.


Toxins

Cucurbitin is an
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha a ...
and a
carboxy In organic chemistry, a carboxylic acid is an organic acid that contains a carboxyl group () attached to an R-group. The general formula of a carboxylic acid is or , with R referring to the alkyl, alkenyl, aryl, or other group. Carboxylic ...
pyrrolidine that is found in raw ''Cucurbita'' seeds. It retards the development of parasitic flukes when administered to infected host mice, although the effect is only seen if administration begins immediately after infection. Cucurmosin is a
ribosome inactivating protein A ribosome-inactivating protein (RIP) is a protein synthesis inhibitor that acts at the eukaryotic ribosome. This protein family describes a large family of such proteins that work by acting as rRNA N-glycosylase (EC 3.2.2.22). They inactivate ...
found in the flesh and seed of ''Cucurbita'', notably ''
Cucurbita moschata ''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 ...
''. Cucurmosin is more toxic to cancer cells than healthy cells.
Cucurbitacin Cucurbitacin is a class of biochemical compounds that some plants – notably members of the pumpkin and gourd family, Cucurbitaceae – produce and which function as a defence against herbivores. Cucurbitacins are chemically classified as t ...
is a plant steroid present in wild ''Cucurbita'' and in each member of the family ''Cucurbitaceae''. Poisonous to mammals, 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. Ingesting too much cucurbitacin can cause stomach cramps, diarrhea and even collapse. 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. While the process of domestication has largely removed the bitterness from cultivated varieties, there are occasional reports of cucurbitacin causing illness in humans. Cucurbitacin is also used as a lure in insect traps.


Pests and diseases

''Cucurbita'' species are used as food plants by the
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
e of some Lepidoptera species, including the
cabbage moth The cabbage moth (''Mamestra brassicae'') is primarily known as a pest that is responsible for severe crop damage of a wide variety of plant species. The common name, cabbage moth, is a misnomer as the species feeds on many fruits, vegetables, ...
(''Mamestra brassicae''), ''
Hypercompe indecisa ''Hypercompe indecisa'' is a moth of the family Erebidae first described by Francis Walker in 1855. It is found in Argentina and Uruguay. Larvae have been recorded feeding on ''Beta'', ''Brassica'', ''Citrus'', ''Cucurbita'', ''Datura'', ''Dio ...
'', and the
turnip moth ''Agrotis segetum'', sometimes known as the turnip moth, is a moth of the family Noctuidae. The species was first described by Michael Denis and Ignaz Schiffermüller in 1775. It is a common European species and it is found in Africa and across ...
(''Agrotis segetum''). ''Cucurbita'' can be susceptible to the pest ''Bemisia argentifolii'' (
silverleaf whitefly The silverleaf whitefly (''Bemisia tabaci'', also informally referred to as the sweet potato whitefly) is one of several species of whitefly that are currently important agricultural pests. A review in 2011 concluded that the silverleaf whitefly ...
) as well as aphids (''
Aphididae The Aphididae are a very large insect family in the aphid superfamily ( Aphidoidea), of the order Hemiptera. These insects suck the sap from plant leaves. Several thousand species are placed in this family, many of which are considered plant/crop ...
''),
cucumber beetle Cucumber beetle is a common name given to members of two genera of beetles, ''Diabrotica'' and ''Acalymma'', both in the family Chrysomelidae. The adults can be found on cucurbits such as cucumbers and a variety of other plants. Many are no ...
s (''Acalymma vittatum'' and '' Diabrotica undecimpunctata howardi''), squash bug (''
Anasa tristis ''Anasa tristis'' is a species of bug in the family Coreidae. It is a major pest of squash and pumpkins, found throughout North America, and is a vector of the cucurbit yellow vine disease bacterium. These bugs can emit an unpleasant odor ...
''), the
squash vine borer The squash vine borer (''Melittia cucurbitae'') is a diurnal species of sesiid moth. The moth is often mistaken for a bee or wasp because of its movements, and the bright orange hind leg scales. The females typically lay their eggs at the base ...
(''Melittia cucurbitae''), and the two-spotted spidermite (''
Tetranychus urticae ''Tetranychus urticae'' (common names include red spider mite and two-spotted spider mite) is a species of plant-feeding mite generally considered to be a pest. It is the most widely known member of the family Tetranychidae or spider mites. It ...
''). The squash bug causes major damage to plants because of its very toxic saliva. The
red pumpkin beetle ''Aulacophora foveicollis'', the red pumpkin beetle, is a species of beetle in the family Chrysomelidae. It is a foliar pest of members of the Cucurbitaceae, particularly the pumpkin. It is also a pest of millets in India. Description The adult ...
(''Aulacophora foveicollis'') is a serious pest of cucurbits, especially the pumpkin, which it can defoliate.
Cucurbits The Cucurbitaceae, also called cucurbits or the gourd family, are a plant family consisting of about 965 species in around 95 genera, of which the most important to humans are: *''Cucurbita'' – squash, pumpkin, zucchini, some gourds *''Lagena ...
are susceptible to diseases such as
bacterial wilt Bacterial wilt is a complex of diseases that occur in plants such as Cucurbitaceae and Solanaceae (tomato, common bean, etc.) and are caused by the pathogens ''Erwinia tracheiphila'', a gram-negative bacterium, or '' Curtobacterium flaccumfacien ...
(''Erwinia tracheiphila''), anthracnose (''
Colletotrichum ''Colletotrichum'' (sexual stage: ''Glomerella'') is a genus of fungi that are symbionts to plants as endophytes (living within the plant) or phytopathogens. Many of the species in this genus are plant pathogens, but some species may have a mutu ...
'' spp.), fusarium wilt (''
Fusarium ''Fusarium'' is a large genus of filamentous fungi, part of a group often referred to as hyphomycetes, widely distributed in soil and associated with plants. Most species are harmless saprobes, and are relatively abundant members of the soil mi ...
'' spp.), phytophthora blight (''
Phytophthora ''Phytophthora'' (from Greek (''phytón''), "plant" and (), "destruction"; "the plant-destroyer") is a genus of plant-damaging oomycetes (water molds), whose member species are capable of causing enormous economic losses on crops worldwide, a ...
'' spp. water molds), and powdery mildew (''
Erysiphe ''Erysiphe'' is a genus of fungi in the family Erysiphaceae. Many of the species in this genus are plant pathogens which cause powdery mildew. Species This genus includes, but is not limited to the following species: * ''Erysiphe alphitoides'' ...
'' spp.). Defensive responses to viral, fungal, and bacterial leaf
pathogen In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a germ ...
s do not involve cucurbitacin. Species in the genus ''Cucurbita'' are susceptible to some types of
mosaic virus A mosaic virus is any virus that causes infected plant foliage to have a mottled appearance. Such viruses come from a variety of unrelated lineages and consequently there is no taxon that unites all mosaic viruses. All the symptoms of each virus ...
including:
cucumber mosaic virus ''Cucumber mosaic virus'' (CMV) is a plant pathogenic virus in the family ''Bromoviridae''. This virus has a worldwide distribution and a very wide host range, having the reputation of the widest host range of any known plant virus. It can be tra ...
(CMV),
papaya ringspot virus ''Papaya ringspot virus'' (PRSV) is a pathogenic plant virus in the genus ''Potyvirus'' and the virus family ''Potyviridae'' which primarily infects the papaya tree. The virus is a non-enveloped, flexuous rod-shaped particle that is between 760& ...
-cucurbit strain (PRSV), squash mosaic virus (SqMV),
tobacco ringspot virus ''Tobacco ringspot virus'' (TRSV) is a plant pathogenic virus in the plant virus family ''Secoviridae''. It is the type species of the genus ''Nepovirus''. Nepoviruses are transmitted between plants by nematodes, thrips, mites, grasshoppers, and ...
(TRSV), watermelon mosaic virus (WMV), and
zucchini yellow mosaic virus ''Zucchini yellow mosaic virus'' (ZYMV) is an aphid-borne potyvirus, regarded as a major pathogen of cucurbits in most regions of the world where these crops are cultivated. ZYMV affects all cucurbits including pumpkins, squashes, vegetable m ...
(ZYMV). PRSV is the only one of these viruses that does not affect all cucurbits. SqMV and CMV are the most common viruses among cucurbits. 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.


Human culture


Culinary uses

Long before European contact, ''Cucurbita'' had been a major food source for the native peoples of the Americas, and the species became an important food for European settlers, including the Pilgrims, even featuring at the first
Thanksgiving Thanksgiving is a national holiday celebrated on various dates in the United States, Canada, Grenada, Saint Lucia, Liberia, and unofficially in countries like Brazil and Philippines. It is also observed in the Netherlander town of Leiden ...
. Commercially produced pumpkin commonly used in
pumpkin pie Pumpkin pie is a dessert pie with a spiced, pumpkin-based custard filling. The pumpkin and pumpkin pie are both a symbol of harvest time, and pumpkin pie is generally eaten during the fall and early winter. In the United States and Canada it is u ...
is most often varieties of ''C. moschata'';
Libby's Libby's (Libby, McNeill & Libby) was an American company that produced canned food and beverages. The firm was established in 1869 in Chicago, Illinois. The Libby's trademark is currently owned by Libby's Brand Holding based in Geneva, Switzerlan ...
, by far the largest producer of processed pumpkin, uses a proprietary strain of the Dickinson pumpkin variety of ''C. moschata'' for its canned pumpkin. Other foods that can be made using members of this genus include biscuits, bread, cheesecake, desserts, donuts,
granola Granola is a breakfast and snack food consisting of rolled oats, nuts, honey or other sweeteners such as brown sugar, and sometimes puffed rice, that is usually baked until crisp, toasted and golden brown. The mixture is stirred while baking ...
, ice cream, lasagna dishes, pancakes, pudding, pumpkin butter, salads, soups, and stuffing.
Squash soup Pumpkin soup is a usually 'bound' (thick) soup made from a purée of pumpkin. It is made by combining the meat of a blended pumpkin with broth or stock. It can be served hot or cold, and is a common Thanksgiving dish in the United States. Var ...
is a dish in African cuisine. The
xerophytic A xerophyte (from Greek ξηρός ''xeros'' 'dry' + φυτόν ''phuton'' 'plant') is a species of plant that has adaptations to survive in an environment with little liquid water, such as a desert such as the Sahara or places in the Alps or t ...
species are proving useful in the search for nutritious foods that grow well in arid regions. ''C. ficifolia'' is used to make soft and mildly alcoholic drinks. In India, squashes (''ghiya'') are cooked with seafood such as prawns. In France, marrows (''courges'') are traditionally served as a
gratin Gratin () is a culinary technique in which an ingredient is topped with a browned crust, often using breadcrumbs, grated cheese, egg or butter.Courtine, Robert J. (ed.) (2003) ''The Concise Larousse Gastronomique'' London: Hamlyn The term ma ...
, sieved and cooked with butter, milk, and egg, and flavored with salt, pepper, and nutmeg, 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 it, Ligure , population_note = , population_blank1_title = , population_blank1 = , demographics_type1 = , demographics1_footnotes = , demographics1_title1 = , demographics1_info1 = , demographics1_title2 ...
, or ''torta di zucca e riso'' from
Emilia-Romagna egl, Emigliàn (man) egl, Emiglièna (woman) rgn, Rumagnòl (man) rgn, Rumagnòla (woman) it, Emiliano (man) it, Emiliana (woman) or it, Romagnolo (man) it, Romagnola (woman) , population_note = , population_blank1_title ...
, the squashes being made into a pie filling with butter,
ricotta Ricotta ( in Italian) is an Italian whey cheese made from sheep, cow, goat, or Italian water buffalo milk whey left over from the production of other cheeses. Like other whey cheeses, it is made by coagulating the proteins that remain after th ...
,
parmesan Parmesan ( it, Parmigiano Reggiano; ) is an Italian hard, granular cheese produced from cows’ milk and aged at least 12 months. It is named after two of the areas which produce it, the provinces of Parma and Reggio Emilia (''Parmigiano'' ...
, egg, and milk; and as a sauce for pasta in dishes like ''spaghetti alle zucchine'' from Sicily. In Japan, squashes such as small ''C. moschata'' pumpkins (''kabocha'') are eaten boiled with sesame sauce, fried as a
tempura is a typical Japanese dish usually consisting of seafood, meat and vegetables that have been battered and deep fried. The dish was introduced by the Portuguese in Nagasaki through fritter-cooking techniques in the 16th century. The word ...
dish, or made into balls with sweet potato and
Japanese mountain yam ''Dioscorea japonica'', known as East Asian mountain yam, yamaimo, or Japanese mountain yam, is a type of yam (''Dioscorea'') native to Japan (including Ryukyu and Bonin Islands), Korea, China, Taiwan, and Assam. ''Dioscorea japonica'' is used ...
.


Art, music, and literature

Along with maize and beans, squash has been depicted in the art work of the native peoples of the Americas for at least 2,000 years. For example, cucurbits are often represented in Moche ceramics. Though native to the western hemisphere, ''Cucurbita'' began to spread to other parts of the world after
Christopher Columbus Christopher Columbus * lij, Cristoffa C(or)ombo * es, link=no, Cristóbal Colón * pt, Cristóvão Colombo * ca, Cristòfor (or ) * la, Christophorus Columbus. (; born between 25 August and 31 October 1451, died 20 May 1506) was a ...
's arrival in the New World in 1492. 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
festoon A festoon (from French ''feston'', Italian ''festone'', from a Late Latin ''festo'', originally a festal garland, Latin ''festum'', feast) is a wreath or garland hanging from two points, and in architecture typically a carved ornament depict ...
s at
Villa Farnesina The Villa Farnesina is a Renaissance suburban villa in the Via della Lungara, in the district of Trastevere in Rome, central Italy. Description The villa was built for Agostino Chigi, a rich Sienese banker and the treasurer of Pope Julius II. B ...
in Rome. Also, in 2001 depictions of this genus were identified in ''
Grandes Heures of Anne of Brittany The Grandes Heures of Anne of Brittany (''Les Grandes Heures d'Anne de Bretagne'' in French) is a book of hours, commissioned by Anne of Brittany, Queen of France to two kings in succession, and illuminated in Tours or perhaps Paris by Jean B ...
'' (''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. In 1952, Stanley Smith Master, using the pen name Edrich Siebert, wrote "The Marrow Song (Oh what a beauty!)" to a tune in time. It became a popular hit in Australia in 1973, and was revived by
the Wurzels The Wurzels are an English Scrumpy and Western band from Somerset, England, best known for their number one hit " The Combine Harvester" and number three hit "I Am a Cider Drinker" in 1976. They are known for using British West Country phrases ...
in Britain on their 2003 album ''Cutler of the West''. John Greenleaf Whittier wrote a poem entitled ''The Pumpkin'' in 1850. "The Great Pumpkin" is a fictional holiday figure in the comic strip ''
Peanuts ''Peanuts'' is a syndicated daily and Sunday American comic strip written and illustrated by Charles M. Schulz. The strip's original run extended from 1950 to 2000, continuing in reruns afterward. ''Peanuts'' is among the most popular and infl ...
'' by
Charles M. Schulz Charles Monroe "Sparky" Schulz (; November 26, 1922 – February 12, 2000) was an American cartoonist and the creator of the comic strip ''Peanuts'', featuring what are probably his two best-known characters, Charlie Brown and Snoopy. He is wi ...
.


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. Pumpkin is also used in cosmetics.


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 Anthelmintics or antihelminthics are a group of antiparasitic drugs that expel parasitic worms (helminths) and other internal parasites from the body by either stunning or killing them and without causing significant damage to the host. They may ...
for the expulsion of worms. In southeastern Europe, seeds of ''C. pepo'' were used to treat irritable bladder and
benign prostatic hyperplasia Benign prostatic hyperplasia (BPH), also called prostate enlargement, is a noncancerous increase in size of the prostate gland. Symptoms may include frequent urination, trouble starting to urinate, weak stream, inability to urinate, or loss o ...
. In Germany, pumpkin seed is approved for use by the
Commission E The German ''Commission E'' is a scientific advisory board of the Federal Institute for Drugs and Medical Devices formed in 1978. The commission gives scientific expertise for the approval of substances and products previously used in traditional, ...
, which assesses folk and herbal medicine, for irritated bladder conditions and
micturition Urination, also known as micturition, is the release of urine from the urinary bladder through the urethra to the outside of the body. It is the urinary system's form of excretion. It is also known medically as micturition, voiding, uresis, ...
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. The
FDA The United States Food and Drug Administration (FDA or US FDA) is a federal agency of the Department of Health and Human Services. The FDA is responsible for protecting and promoting public health through the control and supervision of food ...
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. In China, ''C. moschata'' seeds were also used in
traditional Chinese medicine Traditional Chinese medicine (TCM) is an alternative medical practice drawn from traditional medicine in China. It has been described as "fraught with pseudoscience", with the majority of its treatments having no logical mechanism of acti ...
for the treatment of the parasitic disease schistosomiasis and for the expulsion of
tape worm Cestoda is a class of parasitic worms in the flatworm phylum (Platyhelminthes). Most of the species—and the best-known—are those in the subclass Eucestoda; they are ribbon-like worms as adults, known as tapeworms. Their bodies consist of m ...
s. In Mexico, herbalists use ''C. ficifolia'' in the belief that it reduces
blood sugar Glycaemia, also known as blood sugar level, blood sugar concentration, or blood glucose level is the measure of glucose concentrated in the blood of humans or other animals. Approximately 4 grams of glucose, a simple sugar, is present in the blo ...
levels.


Festivals

''Cucurbita'' fruits including pumpkins and marrows are celebrated in festivals in countries such as Argentina, Austria, Bolivia, Britain, Canada, Croatia, France, Germany, India, Italy, Japan, Peru, Portugal, Spain, Switzerland, and the United States. Argentina holds an annual nationwide pumpkin festival ''Fiesta Nacional del Zapallo'' ("Squashes and Pumpkins National Festival"), in
Ceres, Santa Fe Ceres is a municipality San Cristóbal Department, in Santa Fe Province, Argentina. The town of Ceres is northwest of the provincial capital Santa Fe and has a population of 16,054 The town is situated from the border with Santiago del Es ...
, on the last day of which a ''Reina Nacional del Zapallo'' ("National Queen of the Pumpkin") is chosen. In Portugal the ''Festival da Abóbora de Lourinhã e Atalaia'' ("Squashes and Pumpkins Festival in
Lourinhã Lourinhã () is a municipality in the District of Lisbon, in the Oeste Subregion of Portugal. The population in 2011 was 25,735, in an area of 147.17 km². The seat of the municipality is the town of Lourinhã, with a population of 8,800 inhab ...
and Atalaia") is held in Lourinhã city, called the ''Capital Nacional da Abóbora'' (the "National Capital of Squashes and Pumpkins"). Ludwigsburg, Germany annually hosts the world's largest pumpkin festival. In Britain a giant marrow (zucchini) weighing was displayed in the
Harrogate Harrogate ( ) is a spa town and the administrative centre of the Borough of Harrogate in North Yorkshire, England. Historically in the West Riding of Yorkshire, the town is a tourist destination and its visitor attractions include its spa w ...
Autumn Flower Show in 2012. In the US,
pumpkin chucking Punkin chunkin, or as it is sometimes called pumpkin chucking, is the sport of hurling a pumpkin solely by mechanical means for distance. The devices used include slingshots, catapults, centrifugals, trebuchets, and pneumatic (air) cannons. P ...
is practiced competitively, with machines such as
trebuchet A trebuchet (french: trébuchet) is a type of catapult that uses a long arm to throw a projectile. It was a common powerful siege engine until the advent of gunpowder. The design of a trebuchet allows it to launch projectiles of greater weight ...
s and air cannons designed to throw intact pumpkins as far as possible. The
Keene Pumpkin Fest The New Hampshire Pumpkin Festival (also known as the Keene Pumpkin Festival from 1991 to 2014, and the Laconia Pumpkin Festival in 2015; often referred to as Pumpkin Fest) is a Pumpkin, pumpkin festival that is held in Laconia, New Hampshire, Lac ...
is held annually in
New Hampshire New Hampshire is a state in the New England region of the northeastern United States. It is bordered by Massachusetts to the south, Vermont to the west, Maine and the Gulf of Maine to the east, and the Canadian province of Quebec to the nor ...
; in 2013 it held the world record for the most
jack-o-lantern A jack-o'-lantern (or jack o'lantern) is a carved lantern, most commonly made from a pumpkin or a root vegetable such as a rutabaga or turnip. Jack-o'-lanterns are associated with the Halloween holiday. Its name comes from the reported phenomen ...
s lit in one place, 30,581 on October 19, 2013. Halloween is widely celebrated with jack-o-lanterns made of large orange pumpkins carved with ghoulish faces and illuminated from inside with candles. The pumpkins used for jack-o-lanterns are ''C. pepo'', not to be confused with the ones typically used for pumpkin pie in the United States, which are ''C. moschata''.
Kew Gardens Kew Gardens is a botanic garden in southwest London that houses the "largest and most diverse botanical and mycological collections in the world". Founded in 1840, from the exotic garden at Kew Park, its living collections include some of the ...
marked Halloween in 2013 with a display of pumpkins, including a towering pyramid made of many varieties of squash, in the Waterlily House during its "IncrEdibles" festival.


See also

*
List of gourds and squashes This list of gourds and squashes provides an alphabetical list of (mostly edible) varieties (cultivars) of the plant genus ''Cucurbita'', commonly called gourds, squashes, pumpkins and zucchinis/courgettes. Common names can differ by location. ...
in the genus ''Cucurbita'' * List of squash and pumpkin dishes


Notes


References


External links

* * * {{Authority control Cucurbitaceae genera Squashes and pumpkins Early agriculture in Mesoamerica Crops originating from indigenous Americans Native American cuisine