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Capsicum
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This article is about the plant genus. For the variety known as "capsicum" in some countries, see Bell pepper. For discussion of culinary uses, see Chili pepper.

Capsicum
Capsicum fruit and longitudinal section
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Asterids
Order: Solanales
Family: Solanaceae
Subfamily: Solanoideae
Tribe: Capsiceae
Genus: Capsicum
L.[1]
Species

See text[2]

Capsicum (/ˈkæpsɪkəm/[3]) is a genus of flowering plants in the nightshade family Solanaceae, native to the Americas, cultivated worldwide for their edible fruit, which are generally known as "peppers" or "capsicum". Chili peppers grow on five species of Capsicum. Sweet or bell peppers and some chili peppers are Capsicum annuum, making it the most cultivated species in the genus.[4]

History

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Capsicum is native to South America and Central America.[5] These plants have been evolving for 17 million years.[6] It was domesticated and cultivated at least since 3000 BC, as evidenced by remains of chili peppers found in pottery from Puebla and Oaxaca.[7]

Etymology and names

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The generic name may come from Latin capsa, meaning 'box', presumably alluding to the pods;[8][9] or possibly from the Greek word κάπτω, kapto, 'to gulp'.[10] The name pepper comes from the similarity of piquance (spiciness or "heat") of the flavor to that of black pepper, Piper nigrum, although there is no botanical relationship with it or with Sichuan pepper. The original term chilli came from the Nahuatl word chīlli, denoting a larger Capsicum variety cultivated at least since 3000 BC.[7] Different varieties were cultivated in South America, where they are known as ajíes (singular ají), from the Quechua term for Capsicum.

The fruit (botanically a berry) of Capsicum plants has a variety of names depending on place and type. The more piquant varieties are called chili peppers, or simply chilis. The large, mild form is called bell pepper, or is named by color (green pepper, green bell pepper, red bell pepper, etc.) in North America. In South Africa and some other countries, it is called sweet pepper. The name is simply pepper in the United Kingdom and Ireland.[11] The name capsicum is used in Australia, India, Malaysia, and New Zealand.[12]

Phylogeny

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Capsicums are solanaceous plants within the tribe Capsiceae, and are closely related to Lycianthes.[13]

A 2020 study using ribosomal DNA provided the following phylogenetic tree. It can be seen that in two of the clades, the species C. frutescens is intermingled with C. eximium in one subclade and C. chinense in another subclade; and that C. chacoense is intermingled with C. baccatum.[14]

Capsicum

C. annuum inc. many varieties

C. frutescens (in part) and C. eximium

C. chinense and C. frutescens (in part)

C. baccatum (in part)

C. chacoense (in part)

C. baccatum (in part)

C. chacoense (in part)

C. pubescens inc. Chile de caballo

C. eximium, a pungent chili pepper with purple flowers

C. lycianthoides

Growing conditions

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Ideal growing conditions for peppers include a sunny position with warm, loamy soil, ideally 21 to 29 °C (70 to 84 °F), that is moist but not waterlogged.[15] Extremely moist soils can cause seedlings to "damp-off" and reduce germination.[citation needed]

The plants will tolerate (but do not like) temperatures down to 12 °C (54 °F) and they are sensitive to cold.[16][17] For flowering, Capsicum is a non-photoperiod-sensitive crop.[18] The flowers can self-pollinate. However, at extremely high temperature, 30 to 38 °C (86 to 100 °F), pollen loses viability, and flowers are much less likely to result in fruit.[19]

Species and varieties

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Main article: List of Capsicum cultivars
Red peppers in Cachi, Argentina air-drying before being processed into powder
An arrangement of chilis, including jalapeno, banana, cayenne, and habanero peppers.

Capsicum consists of 20–27 species,[20] five of which are widely cultivated: C. annuum, C. baccatum, C. chinense, C. frutescens, and C. pubescens.[21] Phylogenetic relationships between species have been investigated using biogeographical,[22] morphological,[23] chemosystematic,[24] hybridization,[25] and genetic[20] data. Fruits of Capsicum can vary tremendously in color, shape, and size both between and within species, which has led to confusion over the relationships among taxa.[26] Chemosystematic studies helped distinguish the difference between varieties and species. For example, C. baccatum var. baccatum had the same flavonoids as C. baccatum var. pendulum, which led researchers to believe the two groups belonged to the same species.[24]

Many varieties of the same species can be used in many different ways; for example, C. annuum includes the "bell pepper" variety, which is sold in both its immature green state and in its red, yellow, or orange ripe state. This same species has other varieties, as well, such as the Anaheim chiles often used for stuffing, the dried ancho (before being dried it is called a poblano) chile used to make chili powder, the mild-to-hot, ripe jalapeno used to make smoked jalapeno, known as chipotle.[27]

Peru is thought to be the country with the highest cultivated Capsicum diversity since varieties of all five domesticates are commonly sold in markets in contrast to other countries. Bolivia is considered to be the country where the largest diversity of wild Capsicum peppers are consumed. Bolivian consumers distinguish two basic forms: ulupicas, species with small round fruits including C. eximium, C. cardenasii, C. eshbaughii, and C. caballeroi landraces; and arivivis, with small elongated fruits including C. baccatum var. baccatum and C. chacoense varieties.[28]

The amount of capsaicin is measured in Scoville heat units (SHU) and this value varies significantly among Capsicum varieties. For example, a typical Bell pepper has a value of zero SHU and a Jalapeño has a value of 4000–8000 SHU. In 2017, the Guinness Book of World Records listed the Carolina Reaper as the world's hottest pepper at 1,641,183 SHU, according to tests conducted by Winthrop University in South Carolina, United States. In 2023, the Guinness Book of Records recognized Pepper X as the world's hottest pepper.[29]

Species list

[edit]

Sources:[30][31]

According to Adepoju et al. (2021), the most commonly occurring Capsicum cultivars in Nigeria (and West Africa) are: C. fructescens var. fructescens L.; C. fructescens var. baccatum (L.) Irish; C. annuum var annuum L; C. annuum var. grossum (L.) Sendtn. and C. chinense Jacq.[32]

Formerly placed here

[edit]
  • Tubocapsicum anomalum (Franch. & Sav.) Makino (as C. anomalum Franch. & Sav.)
  • Vassobia fasciculata (Miers) Hunz. (as C. grandiflorum Kuntze)
  • Witheringia stramoniifolia Kunth (as C. stramoniifolium (Kunth) Kuntze)[2]

Genetics

[edit]

Most Capsicum species are 2n=2x=24. A few of the non-domesticated species are 2n=2x=26.[33] All are diploid. The Capsicum annuum and Capsicum chinense genomes were completed in 2014. The Capsicum annuum genome is approximately 3.48 Gb, making it larger than the human genome. Over 75% of the pepper genome is composed of transposable elements, mostly Gypsy elements, distributed widely throughout the genome. The distribution of transposable elements is inversely correlated with gene density. Pepper is predicted to have 34,903 genes, approximately the same number as both tomato and potato, two related species within the family Solanaceae.[34]

Breeding

[edit]

Many types of peppers have been bred for heat, size, and yield. Along with selection of specific fruit traits such as flavor and color, specific pest, disease and abiotic stress resistances are continually being selected. Breeding occurs in several environments dependent on the use of the final variety including but not limited to: conventional, organic, hydroponic, green house and shade house production environments.

Several breeding programs are being conducted by corporations and universities. In the United States, New Mexico State University has released several varieties in the last few years.[35] Cornell University has worked to develop regionally adapted varieties that work better in cooler, damper climates. Other universities such as UC Davis, University of Wisconsin-Madison, and Oregon State University have smaller breeding programs. Many vegetable seed companies breed different types of peppers as well.

Capsaicin

[edit]
Further information: Capsaicin
Molecular structure of capsaicin

The fruit of most species of Capsicum contains capsaicin (methyl-n-vanillyl nonenamide), a lipophilic chemical that can produce a burning sensation (pungency or spiciness) in the mouth of the eater. Most mammals find this unpleasant, whereas birds are unaffected.[36][37] The secretion of capsaicin protects the fruit from consumption by insects.[38]

Capsaicin is present in large quantities in the placental tissue (which holds the seeds), the internal membranes, and to a lesser extent, the other fleshy parts of the fruits of plants in this genus. The seeds themselves do not produce any capsaicin, although the highest concentration of capsaicin can be found in the white pith around the seeds.[39] Most of the capsaicin in a pungent (hot) pepper is concentrated in blisters on the epidermis of the interior ribs (septa) that divide the chambers, or locules, of the fruit to which the seeds are attached.[40]

A study on capsaicin production in fruits of C. chinense showed that capsaicinoids are produced only in the epidermal cells of the interlocular septa of pungent fruits, that blister formation only occurs as a result of capsaicinoid accumulation, and that pungency and blister formation are controlled by a single locus, Pun1, for which there exist at least two recessive alleles that result in non-pungency of C. chinense fruits.[41]

The amount of capsaicin per fruit is highly variable, depending on genetics and the environment. The most recognized Capsicum without capsaicin is the bell pepper,[42] a cultivar of Capsicum annuum, which has a zero rating on the Scoville scale. The lack of capsaicin in bell peppers is due to a recessive gene that eliminates capsaicin and, consequently, the hot taste usually associated with the rest of the genus Capsicum.[43] There are also other peppers without capsaicin, mostly within the Capsicum annuum species, such as the cultivars Giant Marconi,[44] Yummy Sweets,[45] Jimmy Nardello,[46] and Italian Frying peppers (Cubanelles).[47]

Chili peppers are important in the medicine of Indigenous peoples, and capsaicin is used in modern medicine mainly in topical medications as a circulatory stimulant and analgesic. In more recent times, an aerosol extract of capsaicin, known as capsicum or pepper spray, has become used by law enforcement as a nonlethal means of incapacitating a person, and in a more widely dispersed form for riot control, or by individuals for personal defense. Capsaicin in vegetable oils or horticultural products can be used in gardening as a natural insecticide.[48]

Cuisine

[edit]
Chili peppers

Capsicum fruits can be eaten raw or cooked. Those used in cooking are generally varieties of the C. annuum and C. frutescens species, though a few others are used, as well. They are suitable for stuffing with fillings such as cheese, meat, or rice.

They are frequently used both chopped and raw in salads, or cooked in stir-fries or other mixed dishes. They can be sliced into strips and fried, roasted whole or in pieces, or chopped and incorporated into salsas or other sauces, of which they are often a main ingredient.

They can be preserved in the form of a jam,[49] or by drying, pickling, or freezing. Dried Capsicum may be reconstituted whole, or processed into flakes or powders. Pickled or marinated Capsicum are frequently added to sandwiches or salads. Frozen Capsicum are used in stews, soups, and salsas. Extracts can be made and incorporated into hot sauces.

The Spanish conquistadores soon became aware of their culinary properties, and brought them back to Europe, together with cocoa, potatoes, sweet potatoes, tobacco, maize, beans, and turkeys. They also brought it to the Spanish Philippines colonies, whence it spread to Asia. The Portuguese brought them to their African and Asiatic possessions such as India. All varieties were appreciated but the hot ones were particularly appreciated, because they could enliven an otherwise monotonous diet during times of dietary restriction, such as during religious observances.

Spanish cuisine soon benefited from the discovery of chiles in the New World, and it would become very difficult to untangle Spanish cooking from chiles. Ground chiles, or paprika, hot or otherwise, are a key ingredient in chorizo, which is then called picante (if hot chile is added) or dulce (if otherwise). Paprika is an important ingredient in rice dishes, and plays a definitive role in squid Galician style (polbo á feira). Chopped chiles are used in fish or lamb dishes such as ajoarriero or chilindrón. Pisto is a vegetarian stew with chilies and zucchini as main ingredients. They can also be added, finely chopped, to gazpacho as a garnish. In some regions, bacon is salted and dusted in paprika for preservation. Cheese can also be rubbed with paprika to lend it flavor and colour. Dried round chiles called ñoras are used for arroz a banda.

After being introduced by the Portuguese, chile peppers saw widespread adoption throughout South, Southeast, and East Asia, especially in India, Thailand, Vietnam, China, and Korea. Several new cultivars were developed in these countries, and their use in combination with (or as a substitute for) existing 'hot' culinary spices such as black pepper and Sichuan pepper spread rapidly, giving rise to the modern forms a number of staple dishes such as Channa masala, Tom yum, Laziji, and Kimchi. This in turn influenced Anglo-Indian and American Chinese cuisine, most notably with the development of British and American forms of curry powder (based on Indian spice preparations such as garam masala), and dishes such as General Tso's chicken and chicken tikka masala.

Crushed red pepper
Stuffed capsicum with a spicy filling of potatoes, Kolkata, India

According to Richard Pankhurst, C. frutescens (known as barbaré) was so important to the national cuisine of Ethiopia, at least as early as the 19th century, "that it was cultivated extensively in the warmer areas wherever the soil was suitable." Although it was grown in every province, barbaré was especially extensive in Yejju, "which supplied much of Showa, as well as other neighbouring provinces." He mentions the upper Golima River valley as being almost entirely devoted to the cultivation of this plant, where it was harvested year-round.[50]

In 2005, a poll of 2,000 people revealed the capsicum to be Britain's fourth-favourite culinary vegetable.[51]

In Hungary, sweet yellow capsicum – along with tomatoes – is the main ingredient of lecsó.

In Bulgaria, South Serbia, and North Macedonia, capsicum are very popular, too. They can be eaten in salads, like shopska salata; fried and then covered with a dip of tomato paste, onions, garlic, and parsley; or stuffed with a variety of products, such as minced meat and rice, beans, or cottage cheese and eggs. Capsicum are the main ingredient in the traditional tomato and capsicum dip lyutenitsa and ajvar. They are in the base of different kinds of pickled vegetables dishes, turshiya.

Capsicum is also used widely in Italian cuisine, and the hot species are used all around the southern part of Italy as a common spice (sometimes served with olive oil). Capsicums are used in many dishes; they can be cooked by themselves in a variety of ways (roasted, fried, deep-fried) and are a fundamental ingredient for some delicatessen specialities, such as nduja.

Capsicums are also used extensively in Sri Lanka cuisine as side dishes.[52]

The Maya and Aztec people of Mesoamerica used Capsicum fruit in cocoa drinks as a flavouring.[53]

In New Mexico, there is a Capsicum annuum cultivar group called the New Mexico chile which is a mainstay of the state's New Mexican cuisine.

GRAS

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Only Capsicum frutescens L. and Capsicum annuum L. are Generally recognized as safe.[54][55]

Synonyms and common names

[edit]
Capsicum annuum cultivars

The name given to the Capsicum fruits varies between English-speaking countries.

In Australia, New Zealand and Indian English, heatless varieties are called "capsicums", while hot ones are called "chilli"/"chillies" (double L). The term "bell peppers" is never used, although in Australia C. annuum and other varieties which have a bell shape and are fairly hot, are often called "bell chillies".

In Canada, Ireland, South Africa and the United Kingdom, the heatless varieties are known simply as "peppers" (or more specifically "green peppers", "red peppers", etc.), while the hot ones are "chilli"/"chillies" (double L) or "chilli peppers".

In the United States, the common heatless varieties are referred to as "bell peppers", "sweet peppers", "red/green/etc. peppers", or simply "peppers", while the hot varieties are collectively called "chile"/"chiles", "chili"/"chilies", or "chili"/"chile peppers" (one L only), "hot peppers", or named as a specific variety (e.g., banana pepper).

The ground spice made from hot capsicums is called paprika in both U.S. English and Commonwealth English. The name derives from Central Europe; in Polish and in Hungarian, the term papryka and paprika (respectively) is used for all kinds of capsicums and the spice made from them.

Pictures of common cultivars

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See also

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References

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[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Capsicum

View on Grokipedia
from Grokipedia
Capsicum is a of flowering in the nightshade family (), native to Central and , consisting of approximately 25–35 of mostly herbaceous perennials or shrubs that are typically grown as annuals in temperate regions. These are characterized by their alternate, simple leaves, small white to purplish flowers, and berry-like fruits that vary in shape, size, color, and , with the heat in spicy varieties derived from capsaicinoids such as . The originated in the Andean region of northwestern , particularly around and , before diversifying across the . Five species of Capsicum have been domesticated, forming the basis of global pepper cultivation: C. annuum (including bell peppers, jalapeños, and ), C. chinense (habaneros), C. baccatum (aji peppers), C. frutescens (), and C. pubescens (rocoto). occurred independently in multiple regions, with archaeological evidence indicating that C. annuum was cultivated in central-eastern as early as 6000 years ago, while other species were domesticated in . Following Christopher Columbus's voyages in , peppers were introduced to and subsequently spread worldwide through trade, becoming integral to cuisines across , , and beyond. As of 2023, Capsicum species are among the most economically important vegetable crops globally, with annual production of approximately 37 million metric tons, primarily of C. annuum, used fresh, dried, or processed into spices, sauces, and condiments. Their fruits are valued not only for flavor but also for nutritional content, including high levels of vitamins A and C, antioxidants like and , and bioactive compounds with potential medicinal benefits, such as anti-inflammatory and analgesic properties from . Ornamental varieties are also popular in gardening, while wild species contribute to breeding programs for disease resistance and flavor diversity.

Introduction and Overview

Definition and General Characteristics

Capsicum is a of flowering belonging to the family , encompassing approximately 25–35 species, the majority of which are native to the tropical and temperate regions of southern , , and . This is characterized by its diversity in form and adaptation, with typically functioning as herbaceous perennials or shrubs in their native habitats. Morphologically, Capsicum plants range in height from 0.5 to 4 meters, forming upright or bushy structures with simple, ovate to lanceolate leaves that are usually dark green and smooth-edged. The flowers are small, hermaphroditic, and typically white or purplish, measuring 9–15 mm in diameter with a campanulate or rotate corolla. Fruits are botanically classified as berries, exhibiting a wide variation in shape—from spherical to conical or elongated—and color, maturing from green through shades of yellow, orange, or red. A key distinction within the lies between non-pungent () and pungent (hot) varieties, determined by the presence or absence of capsaicinoids, a group of alkaloids primarily concentrated in the placental tissue of the fruit. Pungent types produce these compounds, which impart the characteristic sensation, while types lack them due to recessive alleles at the Pun1 locus. The basic life cycle of Capsicum plants is influenced by : they behave as annuals in temperate regions but as perennials in tropical environments, allowing for multi-year productivity under suitable conditions. Fruit development occurs following flowering, with berries forming through predominantly facilitated by the flowers' structure, although via can contribute up to 90% in some cases.

Economic and Cultural Significance

Capsicum ranks among the world's most important vegetable crops, with global production of green chillies and peppers reaching approximately 41 million metric tons in 2023, cultivated across more than 2 million hectares. China dominates as the leading producer, accounting for over 20.5 million tons that year, followed by Mexico and Indonesia as key contributors. Including dry varieties, total annual output exceeds 40 million tons in recent years, underscoring its scale in global agriculture. Economically, capsicum drives substantial value in international trade, with the global market valued at USD 21.8 billion in and projected to reach USD 30.4 billion by 2030, fueled by demand in fresh produce, processing, and value-added foods like sauces and snacks. It bolsters rural economies in producing nations through exports, where , , and the emerge as top shippers of fresh varieties, while and lead in dried forms for the sector. This not only generates but also supports agro-processing industries, employing millions in harvesting, packaging, and distribution. Culturally, capsicum holds profound significance in diverse societies, symbolizing heat, vitality, and heritage. In , chilies feature prominently in rituals and festivals, such as the , where they are incorporated into traditional offerings and foods to honor ancestors and ward off evil spirits. In , chili peppers are integral to regional identities, infusing everyday cuisine with bold flavors and appearing in ceremonial dishes that reflect communal bonds and agricultural traditions. These roles extend to indigenous practices across and , where capsicum serves as a bridge between sustenance and spirituality. Contemporary trends highlight capsicum's adaptability to modern agriculture, with rising adoption of organic and hybrid varieties enhancing export competitiveness through improved yields and resilience to pests. Hybrid cultivars, in particular, offer higher productivity in organic systems, addressing demands in markets like and . Furthermore, in developing regions, capsicum cultivation aids by providing a versatile, nutrient-dense crop that thrives in varied climates and supports smallholder farmers against climate variability.

History and Etymology

Historical Origins and Domestication

The genus Capsicum, native to the , originated in regions spanning from central to northern , with archaeological evidence indicating early human use and dating back approximately 6,000 years. The earliest substantiated records come from the Tehuacán Valley in , where remains of were found in preceramic contexts around 5,600–6,400 years , suggesting initial gathering and eventual cultivation by local indigenous populations. Similarly, was domesticated in the of , with evidence of use emerging around 4,000 years ago, though direct archaeological finds are sparser compared to C. annuum. was domesticated in the Andean region of and around 6,800 years ago, C. frutescens in the approximately 7,400 years ago, and C. pubescens in the highland about 7,100 years ago. These domestication events marked a shift from wild varieties to cultivated forms selected for larger fruits and reduced , facilitated by early agricultural societies in and the . Prior to European contact, indigenous groups such as the Aztecs and Mayans integrated Capsicum species extensively into their societies across Mesoamerica, utilizing them not only as a staple food for flavoring dishes like stews and sauces but also for medicinal purposes. Aztec codices and Mayan texts describe peppers as remedies for ailments including asthma, coughs, toothaches, and digestive issues, often applied topically or ingested in preparations. Their cultivation alongside maize, beans, and squash in milpa systems underscored their role in sustainable agriculture. Chemical analyses of residues from sites like Chiapas de Corzo confirm Capsicum presence from 400 BCE to 300 CE, highlighting consistent pre-Columbian reliance on these plants for multifaceted utility. The global dissemination of Capsicum accelerated during the following Christopher Columbus's second voyage in 1493, when he returned to with samples of chili peppers from the , mistaking them for the valuable Asian spice. Spanish and Portuguese traders rapidly propagated the plants through their colonial networks, introducing them to by the early and subsequently to via routes and to through Indian Ocean ports. By the mid-16th century, Capsicum had taken root in and African coastal regions, adapting to new climates and integrating into local cuisines. In 1753, formalized the genus in his , classifying key species like C. annuum and C. frutescens based on morphological traits observed in European herbaria. Further advancement occurred in the , when hybridization efforts in the United States and focused on developing milder, larger-fruited varieties suitable for market production, laying groundwork for modern breeding.

Etymology and Common Names

The genus name Capsicum was formally established by Carl Linnaeus in his 1753 publication Species Plantarum, where he classified several pepper species under this term, building on earlier usage by Joseph Pitton de Tournefort in 1719. The etymology is debated, with one prevailing theory deriving it from the Latin capsa, meaning "box" or "case," in reference to the box-like shape of the fruits. An alternative explanation links it to the Greek verb kapto (κάπτω), meaning "to bite," alluding to the pungent, biting sensation produced by many species due to capsaicinoids. Common names for plants in the Capsicum genus vary widely by region, often reflecting linguistic influences from indigenous languages, colonial encounters, and analogies to familiar spices. In English-speaking regions, the fruits are generally called "peppers," a term adopted by in 1493 to describe their resemblance in flavor and use to the unrelated (Piper nigrum), despite their distinct botanical origins in the . The word "chili" (or variants like "chile" and "chilli") originates from the term chīlli, used by indigenous Mesoamerican peoples to denote the pod-like fruits, and entered European languages via Spanish after the . In Spanish, "pimiento" derives from pigmentum ("" or "coloring"), highlighting the vibrant hues of the pods, and is commonly applied to milder varieties. Hindi speakers use "mirch," borrowed from marīca (originally referring to ), which was extended to Capsicum species upon their introduction to the . In , particularly among Quechua and Aymara communities, "aji" stems from the word aší (via indigenous languages), denoting hot peppers and persisting in modern Spanish as a regional synonym. Specific variations in naming often distinguish between pungent and non-pungent types or highlight cultivars. Sweet, bell-shaped varieties of Capsicum annuum are known as "bell peppers" in North American English, emphasizing their shape and lack of heat, while in Australia, New Zealand, and parts of India, the broader term "capsicum" is reserved for these mild fruits to differentiate them from spicier "chilies." Named cultivars like "jalapeño," from the Nahuatl xālapa (referring to the Mexican city of Jalapa), denote a specific medium-heat C. annuum pod used fresh or pickled. Following the in the 16th century, Capsicum species spread rapidly from the to and via and Spanish traders, leading to the of names into local and languages. In parts of , such as , terms like "pilipili" evolved from pimenta ("pepper"), applied to both hot chilies and milder types in trade contexts. In , introductions prompted borrowings like Indonesian "cabe" (from pimenta) and the extension of existing spice words, such as "mirch" in , integrating the plants into diverse culinary traditions within decades.

Taxonomy and Classification

Phylogeny and Evolutionary Relationships

The genus Capsicum is placed within the Solanaceae family, alongside economically important crops like potatoes (Solanum tuberosum) and tomatoes (S. lycopersicum), and is more closely related to Physalis (ground cherry) than to Solanum. Recent telomere-to-telomere genome assemblies indicate that Capsicum diverged from Physalis approximately 17 million years ago and from Solanum around 19 million years ago, during the mid-Miocene epoch. Earlier whole-genome comparisons had estimated a deeper split from tomato and potato at about 36 million years ago, but updated phylogenetic analyses using thousands of orthologous genes support the more recent divergence within Solanaceae. Capsicum forms a monophyletic , with Lycianthes as its closest , originating in the of western (, , ) before expanding clockwise around the to central and southeastern , and northward to . Phylogenetic reconstructions, based on chloroplast DNA markers such as matK, psbA-trnH, and the atpB-rbcL spacer, along with nuclear genes like waxy, delineate 9 to 11 major , including Andean, , Flexuosum, , Bolivian, Pubescens, Tovarii, Baccatum, and Annuum. The five main domesticated —C. annuum, C. chinense, C. frutescens, C. baccatum, and C. pubescens—cluster within derived (Annuum, Baccatum, and Pubescens), distinct from their wild ancestors, with diversification accelerating from the (~5 million years ago) to the Pleistocene (~0.5–1 million years ago) amid geological and climatic shifts. A key evolutionary adaptation in Capsicum is the development of capsaicinoids, pungent compounds that serve as a , deterring mammals—which crush seeds during digestion and reduce viability—while sparing birds, which disperse intact seeds over long distances. This directed deterrence likely arose after the genus's divergence from , with capsaicin synthase genes duplicating in around 71 million years ago and the full biosynthetic pathway evolving between 13.4 and 5 million years ago, enabling tissue-specific production in fruit placentas. Interspecific hybridization faces post-fertilization barriers, such as endosperm breakdown, but limited occurs among wild populations and between wild and domesticated forms, facilitated by shared habitats and pollinators. Post-2020 genomic sequencing efforts, including gapless assemblies of C. annuum and wild relatives like C. rhomboideum, have illuminated polyploidy-like duplications in biosynthetic pathways and from wild species into domesticated lineages, such as 1.71% from the Baccatum to the Annuum , enriching traits like disease resistance. A graph of 500 accessions across five domesticated species and wild relatives identified over 100 million SNPs, revealing adaptive on chromosomes 6, 7, and 12, while confirming the diploid nature (2n=24) of most Capsicum taxa without recent whole-genome duplications. These studies underscore the role of wild pools in diversification and highlight Central Andean hotspots as centers of ongoing .

Species and Varieties

The Capsicum comprises 43 accepted and five varieties, as recognized in a comprehensive 2022 taxonomic based on morphological, anatomical, karyological, and phylogenetic data. Of these, five are domesticated, originating from wild ancestors in the , while the remaining are wild and contribute to the 's . The domesticated include C. annuum L., which encompasses a wide array of mild and hot peppers such as bell peppers, jalapeños, and ; C. chinense Jacq., known for intensely pungent varieties like ; C. baccatum L., featuring fruity ají-type peppers; C. frutescens L., including and African bird's eye chilies; and C. pubescens (Ruiz & Pav.) Kunth, represented by rocoto peppers with black seeds and flowers. Wild , often shrubby or prostrate, include C. flexuosum Hunz., C. lanceolatum Hunz., C. benoistii Hunz., C. caatingae Barboza & , C. caballeroi M.Nee, C. buforum Hunz., C. campylophorum Hunz., C. cardenasii Heiser & P.M.Sm., C. ceratocalyx (Miers) Kuntze, C. chacoense Hunz., C. coccineum (Rusby) Hunz., C. corallinum Hunz., C. dimorphum Hunz., C. dusenii Hunz., C. ecuadorense Barboza, C. eximium Hunz., C. friburgense Hunz., C. geminifolium Dunal, C. globosum Hunz., C. huarochiriense Hunz., C. huntianum Hunz., C. lycianthoides Bitter, C. mirum Barboza, C. muticum (Sendtn.) Barboza, C. parvifolium Hunz., C. pachyceras Hunz., C. praetermissum Hunz., C. ramiflorum Vell., C. ramosii Hunz., C. recurvatum Hunz., C. schottianum Sendtn., C. scoriosperma Hunz., C. selvae Hunz., C. somniferum Barboza, C. tovarii Eshbaugh, and C. villosum Kunth, among others described in recent revisions. These species exhibit significant varietal diversity, particularly within the domesticated taxa, with approximately 50,000 named cultivars worldwide, varying in fruit shape (e.g., conical, blocky, elongated), color (green to red, yellow, purple), and pungency. Pungency, determined by capsaicinoid concentration, is quantified using the Scoville scale in heat units (SHU), spanning from 0 SHU for sweet bell peppers (C. annuum) to over 1,000,000 SHU for ultra-hot varieties like some C. chinense cultivars. Representative examples include the mild C. annuum 'California Wonder' bell pepper, medium-heat jalapeño (C. annuum), hot cayenne (C. annuum), fruity aji amarillo (C. baccatum), and pungent rocoto (C. pubescens), selected for culinary, ornamental, or ornamental uses based on these traits. Taxonomic revisions in the genus have incorporated DNA-based phylogenies from 2020s studies, refining species boundaries and splitting subgroups within clades; for instance, genome-wide RAD-seq data updated the evolutionary hypothesis for nearly all accepted , confirming and relationships among wild taxa. Some taxa formerly placed in Capsicum have been reclassified into related genera like Lycianthes or Vassobia following molecular evidence distinguishing floral and fruit characteristics; examples include Tubocapsicum anomalum (formerly C. anomalum) and Vassobia fasciculata (formerly C. grandiflorum). hotspots for Capsicum, particularly the C. baccatum complex, are concentrated in the Andean regions of and , where wild and domesticated forms co-occur, supporting high essential for conservation.

Cultivation and Production

Growing Conditions and Requirements

Capsicum , native to tropical and subtropical regions of the , thrive in warm climates with optimal daytime temperatures ranging from 20°C to 30°C and nighttime temperatures between 15°C and 18°C. These are highly sensitive to frost, with exposure below 0°C causing severe damage or death, and prolonged temperatures below 10°C inhibiting growth. Most varieties exhibit day-length neutrality, allowing flowering and fruit set regardless of photoperiod length, which facilitates cultivation in diverse latitudes. For soil, Capsicum prefers well-drained, fertile loamy soils with a range of 6.0 to 7.0 to support uptake and development. Sandy or textures are ideal, as heavy clay soils can lead to waterlogging and . To ensure optimal nutrient availability, growers should monitor plants for visual symptoms of deficiencies. Nitrogen deficiency typically presents as yellowing of older leaves; phosphorus deficiency leads to weak stems and stunted growth; potassium deficiency causes scorched or brown edges on leaf margins; and calcium deficiency manifests as blossom end rot, a dark, sunken lesion at the fruit's blossom end. Both phosphorus (P) and potassium (K) are essential for fruit set and yield in hot peppers (Capsicum spp.), though they play different roles. Phosphorus is critical for early root development, flower formation, and reproductive organ development, directly supporting fruit set; deficiency reduces fruit production. Potassium is vital during flowering, fruit set, and maturation, enhancing carbohydrate accumulation, fruit size, quality, disease resistance, and overall yield; its demand increases significantly in reproductive stages. For optimal results, apply based on soil tests—higher P early (e.g., N-P-K ratio favoring higher P such as 1:2:1), then higher K during fruiting (e.g., 2:1:3)—as excess N can hinder set, while balanced P and K maximize yield. Regular monitoring and adjustments to fertilization based on these symptoms, along with soil testing, can prevent issues and enhance yields. Regarding water requirements, the crop typically needs 500 to 800 mm of irrigation annually, depending on rainfall, , and growth stage, with systems recommended for efficient water delivery and reduced disease risk. Propagation is primarily achieved through seeds, which are sown indoors 6 to 8 weeks before the last frost to ensure vigorous transplants. Germination occurs in 7 to 14 days at soil temperatures of 25°C to 30°C, requiring consistent moisture and light warmth. Transplants are set out after soil warms to at least 15°C, spaced 30 to 60 cm apart in rows 75 to 90 cm wide to optimize airflow and light penetration. Harvesting begins 60 to 90 days after , when fruits reach full size and desired color, with multiple pickings extending the season. Yields for bell peppers typically range from 25 to 50 tons per under optimal conditions, influenced by variety, climate, and management practices.

Breeding, , and Varietal Development

Domestication of Capsicum species has led to significant genetic bottlenecks, reducing variability in cultivated lines compared to their wild progenitors, which limits adaptation to new stresses and environments. Wild relatives, such as and Capsicum chacoense, harbor valuable alleles for traits like disease resistance that have been introgressed into elite varieties through interspecific hybridization to broaden the genetic base. For instance, genes conferring resistance to pathogens like capsici have been successfully transferred from wild species into cultivars via programs. Conventional breeding in Capsicum relies heavily on hybridization to develop F1 hybrids, which provide uniformity in fruit shape, size, and maturity for commercial production. (MAS) has accelerated progress by targeting specific loci for traits like resistance, using molecular markers to select progeny without extensive phenotyping. Recent advances include CRISPR/Cas9 editing, applied post-2020 to knock out susceptibility genes for enhanced resistance to bacterial spot and other diseases in pepper varieties. Breeding programs prioritize traits such as tolerance during reproduction, higher yield under stress, and improved color retention in fruits to extend . has produced super-hot varieties like the in the 2010s, achieved through crosses between landraces to amplify capsaicinoid levels beyond 1.5 million Scoville Heat Units. The sequencing of the genome in 2014 has been pivotal, enabling high-resolution QTL mapping for like fruit size, where major loci on chromosomes 2 and 4 explain significant phenotypic variation. These genomic resources facilitate precise breeding by identifying candidate genes for and editing.

Chemical Composition and Properties

Capsaicinoids and Pungency

Capsaicinoids are a group of alkaloids unique to the Capsicum, responsible for the characteristic or "" in chili peppers. The primary compound, (trans-8-methyl-N-vanillyl-6-nonenamide), is a crystalline, lipophilic, colorless, and odorless substance with the molecular formula C18H27NO3. accounts for approximately 70-90% of total capsaicinoids in most pungent varieties and binds to the receptor in sensory neurons, triggering the burning sensation. Biosynthesis of capsaicin occurs exclusively in the epidermal glandular cells of the placental tissue within the fruit, where precursors from two convergent pathways are combined. The phenylpropanoid pathway derives vanillylamine from phenylalanine through a series of enzymatic steps, including action by phenylalanine ammonia-lyase. Concurrently, the branched-chain fatty acid pathway, starting from valine (or leucine), produces 8-methyl-6-nonenoyl-CoA via enzymes such as branched-chain aminotransferase and acyl-CoA synthetase. These intermediates are then condensed by capsaicin synthase (CS), an acyltransferase, to form capsaicin, with accumulation peaking during fruit maturation. Environmental factors like temperature and nutrient availability, particularly nitrogen sources, influence this process by modulating precursor supply. Related capsaicinoids include , , homocapsaicin, and homodihydrocapsaicin, which share structural similarities but differ in saturation or chain length. , the second most abundant (typically 20-30% of total), lacks the present in and contributes significantly to overall . Homocapsaicin and homodihydrocapsaicin have an extended alkyl chain, comprising 1-10% in various species. The total concentration of these capsaicinoids, rather than any single compound, determines the fruit's level, with variations arising from genetic and environmental factors. Pungency is quantified using the Scoville Heat Units (SHU) scale, developed in 1912 by pharmacologist as an test involving of pepper extract until the heat is undetectable by trained tasters. Originally subjective, the scale has been refined with (HPLC), which directly measures total capsaicinoid concentration in parts per million (ppm); SHU is calculated by multiplying ppm by 16, as pure equates to 16 million SHU. For example, non-pungent bell peppers (C. annuum) register 0 SHU, while highly pungent habanero peppers (C. chinense) range from 100,000 to 350,000 SHU, reflecting ppm levels of 6,250-21,875. This method provides precise, reproducible results for breeding and . In wild Capsicum species, serves key ecological roles, primarily as a . It acts as a deterrent against mammals, which experience intense pain from activation and thus avoid consuming fruits that could damage intact , while birds—lacking this sensitivity—readily eat the peppers and disperse viable via endozoochory. Recent studies confirm 's antimicrobial properties, particularly against fungal pathogens like and species that threaten viability in humid environments; concentrations above 0.1% inhibit germination and mycelial growth. This dual function enhances survival and dispersal in natural habitats.

Nutritional and Phytochemical Profile

Capsicum fruits, primarily from species such as , are characterized by a high water content of approximately 92%, contributing to their low caloric density of 20–40 kcal per 100 g. They provide modest amounts of macronutrients, including about 1 g of protein, 6 g of carbohydrates (with 2–4 g of sugars and 2 g of ), and negligible (0.3 g per 100 g). These values are consistent across varieties, making Capsicum a nutrient-dense, low-energy option. In terms of vitamins, Capsicum is particularly rich in ascorbic acid (), with red varieties containing up to 190 mg per 100 g, exceeding the daily recommended intake for adults in a single serving. Provitamin A , such as beta-carotene, are also prominent, especially in mature red fruits, providing around 3,100 IU of equivalents per 100 g. Other micronutrients include (tocopherols) and like , though in lower concentrations. The profile of Capsicum includes a diverse array of bioactive compounds, notably such as beta-carotene and capsanthin, which impart color and contribute to provitamin A activity. Flavonoids like and polyphenols, including phenolic acids (e.g., ferulic and sinapic acids), are abundant, with total phenolic content ranging from 1–2 mg equivalents per gram in fresh fruits. These compounds underpin the high capacity of Capsicum, often measured by ORAC values of 6,000–11,000 μmol TE per 100 g in sweet varieties, reflecting their potential to neutralize free radicals. Nutritional variations exist between sweet and hot Capsicum types; sweet peppers (e.g., bell varieties) are generally richer in (up to 200 mg/100 g in reds), while hot peppers contain higher levels of capsaicinoids, as detailed in the section on capsaicinoids and . Ripeness also influences composition, with mature red fruits accumulating more and compared to green, immature ones. Recent studies from the have highlighted the presence of phenolics, such as derivatives, in Capsicum extracts, with total phenolic contents supporting roles. Bioavailability of these nutrients can be affected by cooking methods; boiling and steaming reduce vitamin C retention by 25–66% due to leaching into water, whereas stir-frying and roasting preserve more and even enhance accessibility through breakdown. Dry-heat methods like microwaving show minimal impact on total phenolics and antioxidant activity, making them preferable for retaining benefits.

Uses and Applications

Culinary Applications

Capsicum species, commonly known as peppers or chilies, play a central role in global cuisines, valued for their flavor, heat, and color. In cooking, peppers () are a staple in fresh salsas, where they provide pungency and freshness when chopped raw with tomatoes, onions, and cilantro. Indian curries frequently incorporate dried or fresh chilies ( var. longum) to add heat and depth, as seen in spice blends like or dishes such as , where they balance sweetness and acidity in tomato-based gravies. In , gochugaru—coarse powder from sun-dried —is essential for , imparting a fruity spiciness during with and other vegetables. Preparation techniques for Capsicum enhance its versatility in dishes, from fresh applications in salads to cooked forms in stews. peppers over an open flame or in an oven blisters the skin for easy removal, concentrating flavors for use in sauces or stuffed preparations, a method common in Mediterranean and Latin American recipes. involves blanching peppers in with spices, preserving them for extended and adding tanginess to tacos or antipasti. Fermenting peppers in a salt , often lacto-fermentation, develops richness and complex , as in hot sauces or Korean paste. To manage the heat from capsaicinoids, culinary traditions pair Capsicum with dairy like or cheese, or acids such as lime, which neutralize pungency through protein binding or dilution. The U.S. recognizes and its capsaicinoids as (GRAS) for use as spices and flavorings in food, based on historical consumption patterns and safety data. Estimated dietary intake of capsaicinoids from spices in some countries ranges from 25 to 200 mg per person, depending on regional consumption habits, though intakes approaching the upper end may cause adverse gastrointestinal effects in some individuals. In the , Capsicum extracts serve as natural colorants and flavor enhancers, particularly derived from , which imparts red hues and mild heat to sauces, processed meats, and snacks. Recent trends highlight Capsicum in fusion cuisines, blending elements like Korean gochugaru in tacos or Indian chili pastes in Italian pastas, reflecting global flavor experimentation.

Medicinal and Pharmacological Uses

Capsicum, particularly species like Capsicum annuum and Capsicum frutescens, has been employed in traditional medicine systems for its anti-inflammatory properties. In Ayurveda, chili peppers are used to alleviate joint pain and arthritis symptoms, often through topical applications such as oil massages that leverage capsaicin's counter-irritant effects to reduce inflammation. Folk medicine traditions, including those in India and other regions, incorporate Capsicum for its antimicrobial qualities, traditionally as a digestive aid, carminative, and treatment for diarrhea due to its inhibitory effects on bacterial pathogens. In modern , , the primary in Capsicum, is widely utilized for topical pain relief in concentrations of 0.025% to 0.075% in creams, demonstrating efficacy against such as and through desensitization of pain receptors. Clinical trials since 2010 have explored intranasal or topical applications for relief, showing reductions in headache severity and arterial tenderness by modulating activity, though results vary and further studies are needed for broader adoption. Beyond , other Capsicum constituents contribute to therapeutic potential. High levels of in bell peppers support immune function by enhancing activity and acting as an to protect against . such as , , and capsanthin in colored Capsicum varieties promote eye health by accumulating in the to filter blue light and reduce the risk of age-related . Ongoing research highlights capsaicin's anticancer effects, including induction of in various lines like and oral cancers via generation and arrest. Safety considerations are important, as high doses of Capsicum or can cause gastrointestinal irritation, including , , and , particularly in sensitive individuals. This occurs because capsaicin binds to transient receptor potential vanilloid 1 (TRPV1) receptors in the gastrointestinal tract, stimulating strong intestinal contractions that can lead to abdominal cramps and accelerated transit time resulting in diarrhea. It is contraindicated for patients with (GERD) due to exacerbation of esophageal symptoms from its irritant properties.

Conservation and Challenges

Pests, Diseases, and Environmental Issues

Capsicum crops are susceptible to several major insect pests, including aphids (Myzus persicae and other species), thrips (Frankliniella occidentalis and Thrips tabaci), and spider mites (Tetranychus urticae). Aphids feed by sucking sap from leaves and stems, causing curling, stunting, and yellowing, while also serving as vectors for viruses such as cucumber mosaic virus and potato virus Y. Thrips damage results in distorted foliage, silvering, and crinkled leaves, often leading to reduced photosynthesis and fruit quality. Spider mites, particularly the two-spotted variety, produce fine webbing on leaf undersides and cause stippling, bronzing, and premature leaf drop under hot, dry conditions. Diseases pose significant threats to Capsicum production, with viral, fungal, and bacterial pathogens being prominent. (TMV), a , induces mosaic patterns, leaf mottling, and plant stunting; it spreads mechanically through infected tools, hands, or seed, and via products. blight, caused by the Phytophthora capsici, manifests as , stem lesions, and fruit decay, thriving in warm, wet soils; transmission occurs through infested soil, water splash, or infected plant debris, leading to rapid crop losses in poorly drained fields. Bacterial spot, induced by Xanthomonas euvesicatoria, produces water-soaked lesions on leaves and fruit that turn necrotic with yellow halos; it spreads via rain splash, overhead , or contaminated , exacerbated by high . Resistant varieties, such as those with the Bs2 for bacterial spot or Phyto loci for P. capsici, have been developed to mitigate these issues. Environmental challenges further compound production risks for Capsicum. Water stress from or deficit reduces shoot growth, root architecture, and overall vigor, leading to lower set and yield. impairs nutrient and water uptake, causing osmotic stress, reduced content, and stunted development, with even moderate salt levels (e.g., 2-4 dS/m) significantly impacting sensitive varieties. exacerbates these vulnerabilities, with projections indicating yield declines in tropical regions by the 2030s due to rising temperatures, erratic rainfall, and increased , shifting suitable growing zones and heightening pressure. Mitigation strategies emphasize (IPM), including biological controls like predatory mites (Phytoseiulus persimilis) for spider mites and parasitic wasps for , alongside cultural practices such as to break pest cycles and methods to enhance and . For diseases, , applications, and rotation with non-host crops (e.g., cereals for 2-3 years) reduce buildup, while recent gene-editing techniques like / have introduced resistance traits, such as CaMLO2 knockout for broad-spectrum fungal protection, complementing conventional breeding efforts.

Conservation Status and Biodiversity

Many wild species in the genus Capsicum face significant conservation challenges, with assessments indicating that a substantial portion are at risk of . According to preliminary threat evaluations based on modelled distributions across 37 wild taxa, six are potentially critically endangered, three endangered, ten vulnerable, six near threatened, and twelve least concern. For instance, Capsicum lanceolatum, a rare Mesoamerican known from limited natural occurrences, is classified as endangered due to ongoing habitat loss from . hotspots for wild Capsicum are concentrated in the Andean regions, including tropical forests and inter-Andean valleys in countries like , , , and , where approximately 50% of the genus's diversity occurs. These areas harbor high but are under pressure from environmental changes. Biodiversity loss in wild Capsicum populations is primarily driven by and , which have converted native habitats into agricultural lands, as seen in the destruction of sites for C. lanceolatum. Additionally, gene erosion arises from hybridization between domesticated crops and wild relatives, leading to that dilutes genetic distinctiveness in natural populations. In , over 80% of original native vegetation has been lost, exacerbating threats to crop wild relatives like Capsicum, with 35% of assessed taxa threatened according to IUCN criteria. Conservation strategies emphasize both ex situ and approaches to safeguard Capsicum genetic resources. Globally, over 50,000 accessions are conserved in genebanks, with key collections at CGIAR-affiliated centers such as the World Vegetable Center, which holds approximately 9,171 Capsicum accessions. efforts include protected areas in , such as regions supporting wild C. annuum var. glabriusculum, and in , where initiatives focus on endemic like C. caatingae through habitat preservation and on-farm management. These strategies prioritize collecting from underrepresented wild taxa, with 35 of 37 wild targeted for enhanced representation. Recent initiatives in the , including the Global Strategy for the Conservation and Use of Capsicum Genetic Resources, promote the of conserved genes to bolster wild populations by integrating ex situ materials into restoration projects in hotspots. This work aligns with international agrobiodiversity treaties, such as the (CBD), which underscores the sustainable use and conservation of plant genetic resources, and supports related protocols like the for access and benefit-sharing.

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