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4 December 2014

Capsicum baccatum (pepper)

Datasheet Types: Crop, Invasive species, Host plant

Abstract

This datasheet on Capsicum baccatum covers Identity, Overview, Associated Diseases, Pests or Pathogens, Distribution, Dispersal, Biology & Ecology, Environmental Requirements, Impacts, Uses, Management, Genetics and Breeding, Food Quality, Economics, Further Information.

Identity

Preferred Scientific Name
Capsicum baccatum L.
Preferred Common Name
pepper
Other Scientific Names
Capsicum cerasiflorum Link
Capsicum baccatum L. var. pendulum (Willd.) Eshbaugh
Capsicum chamaecerasus Nees
Capsicum ciliare Willd.
Capsicum conicum Vell.
Capsicum microcarpum Cav.
Capsicum microphyllum Dunal
Capsicum pulchellum Salisb.
Capsicum umbilicatum Vell.
International Common Names
English
aji
bishop's-hat
Christmas bell
Peruvian pepper
French
piment chien
Local Common Names
Bolivia
ají
arivivi
Dominican Republic
ají bobito
ají bonito
ají caribe
ají montesino
ají tití
ajicito montesino
Germany
peruanischer Pfeffer
Haiti
piment zouézeau
piment zouézo
pimento z'oiseux
Puerto Rico
ají caballero
ají pico de paloma
Sweden
barpeppar
United States Virgin Islands
wild pepper

Pictures

Capsicum baccatum (pepper, bishop's hat, orchid pepper); fruiting habit. Pali o Waipio Huelo, Maui, Hawaii, USA. September, 2014
Fruiting habit
Capsicum baccatum (pepper, bishop's hat, orchid pepper); fruiting habit. Pali o Waipio Huelo, Maui, Hawaii, USA. September, 2014
©Forest & Kim Starr-2014 - CC BY 3.0
Capsicum baccatum (pepper, bishop's hat, orchid pepper); fruiting habit. Pali o Waipio Huelo, Maui, Hawaii, USA. September, 2014.
Fruiting habit
Capsicum baccatum (pepper, bishop's hat, orchid pepper); fruiting habit. Pali o Waipio Huelo, Maui, Hawaii, USA. September, 2014.
©Forest & Kim Starr-2014 - CC BY 3.0
Capsicum baccatum (pepper, bishop's hat, orchid pepper); close-up of fruits. Pali o Waipio Huelo, Maui, Hawaii, USA. September, 2014.
Fruits
Capsicum baccatum (pepper, bishop's hat, orchid pepper); close-up of fruits. Pali o Waipio Huelo, Maui, Hawaii, USA. September, 2014.
©Forest & Kim Starr-2014 - CC BY 3.0

Overview

A domesticate of the Andean region, Capsicum baccatum, is still little cultivated outside of South America and the most commonly grown species on the continent. It is often called aji, the name also used for other pungent peppers in South America and the West Indies. It can readily be distinguished from the other cultivated peppers by the presence of yellow spots on the corolla. The centre of diversity for C. baccatum is either Bolivia or Peru. Consisting of wild and domesticated genotypes, the species is a valuable source of new genes useful for improving fruit quality and disease resistance in C. annuum sweet bell and hot chilli peppers. The wild forms of C. baccatum have small, erect deciduous fruits, while the domesticated form, recognised as var. pendulum, has larger-sized fruits which are pendant and remain attached to the plant. During domestication, selection has led to larger fruits with increased weight that has resulted in their pendulous nature, and their retention on the plant. As many different pod types (in relation to shape, colour and size) exist in C. baccatum as in C. annuum. The fruits also vary in pungency, from very mild to fiery hot.
Fruits are used extensively in cooking and medicinally, e.g. as a carminative, stimulant, digestive and pain reliever. Usually used fresh or dried, they can also be canned or pickled in brine.
Propagated by seed, peppers grow best on well-drained, sandy or silt-loam soil. Optimal temperatures for fruit production are between 20 and 30°C.

Summary of Invasiveness

C. baccatum is a perennial plant listed as a ‘weed’ in the Global Compendium of Weeds (Randall, 2012), is reported to be invasive to Cuba (Oviedo-Prieto et al., 2012), and is reportedly a weed in Trinidad and Brazil (Holm et al., 1979; Randall, 2012). The species has been cultivated since pre-Columbian times in its native South America and later around the world, for use as a highly popular spice, vegetable, ornamental, and ingredient in such commodities as self defense pepper spray (Basu and De, 2003). The species reproduces by seeds encased in its famous chilli fruits, which are spread primarily by intentional and accidental biotic dispersal agents.

Taxonomic Tree

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Notes on Taxonomy and Nomenclature

The genus Capsicum consists of all the ‘chilli pepper plants’ with 3-5 wild species and over 2000 cultivars (DeWitt and Bosland, 1996; Tewkesbury et al., 2006), and the confusing terminology ‘chilli’ is often used frequently and interchangeably with other names including ‘chile’, ‘aji’, and ‘paprika’ referring to multiple species (Basu and De, 2003). The genus name Capsicum derives from a Greek-based derivative of the latin word ‘kapto’, meaning ‘to bite’, in reference to the heat or pungency of the species’ fruit (Basu and De, 2003), although it has also been speculated to derive from the Latin word ‘capsa’, a box, referring to the shape of the fruit in forms of the typical species (Britton, 1918). The common name ‘chile’ is a variation of ‘chil’, derived from the Nahutal (Aztec) dialect (Basu and De, 2003).
The number of global species within the Capsicum genus has long been subject to debate, but there are presently considered to be five domesticated species of Capsicum, the primary distinguishing characteristics being flower and seed colour, shape of the calyx, number of flowers per node and their orientation; these five species are C. annuum, C. baccatum var. pendulum, C. frutescens, C. chinense, and C. pubescens (DeWitt and Bosland, 1996; Hawkes et al., 1979; Basu and De, 2003).
C. baccatum was first described by Linnaeus in 1753, though this name has also been misapplied to a number of different taxa. The type material of C. baccatum can be matched with material found in the wild today; however, in his original description in the Species Plantarum, Linnaeus failed to mention the presence of white flowers with yellow corolla markings that are apparent in the type material. This led a number of workers to rely entirely on plant habit and fruit shape for their association of living plant material with the original description (Eshbaugh, 1970).
Historically, C. baccatum has been separated into the two species, C. microcarpum and C. pendulum, a separation based primarily on differences in the fruit characters. Today, separation of the wild and cultivated taxa is weak and is maintained primarily through geographic isolation outside the range of wild C. baccatum and by agricultural isolation within the overlap zone of the two varieties (Eshbaugh, 1970).
The species C. baccatum has both a wild form, C. baccatum var. baccatum, and a cultivated subspecies, C. baccatum var. pendulum; the cultivated form is widespread throughout tropical regions in South America, while the wild form is more restricted but ranges from Peru to Brazil (Basu and De, 2003). Eshbaugh (1970) described the qualitative characters of fruit colour, fruit position, and fruit persistence as easily distinguishing the two varieties of C. baccatum. C. baccatum var. baccatum has red, erect, and non-persistent fruits, and C. baccatum var. pendulum has red, orange, yellow, green, or brown fruits which are pendent and persistent. Basu and De (2003) later reported that the two are morphologically indistinguishable, with identical flavonoid and isoenyzme profiles, except for the organ size differences in var. pendulum.
An AFLP study of C. baccatum accessions from South America did not support taxonomic distinction of C. baccatum var. umbilicum from C. baccatum var. pendulum (Albrecht et al., 2012). A clustering analysis of the same data suggested that C. baccatum likely originated in Paraguay. The Plant List (2016) listed only 2 accepted names for infraspecific taxa of C. baccatum: var. pendulum (Willd.) Eshbaugh and var. praetermissum (Heiser & P.G. Sm.) Hunz.

Plant Type

Herbaceous
Perennial
Seed propagated
Shrub

Description

Erect or scrambling, often much branched, perennial herb to sappy shrub, up to 4.5 m (but often less). Branches ± angular, slightly striate, densely pubescent when young to ± glabrous. Leaves usually solitary, rarely 2 appearing together; petiole 0.3–3 cm long; lamina membranous, 1.5–7.5(10) × 0.8–4(4.5) cm, ovate to ovate-lanceolate, base rounded to narrowly cuneate, and often unequal-sided, apex long-acuminate, entire, ± ciliate, with scattered hairs, sometimes only along the nerves, paler and with a few hairs in the axils of the nerves beneath. Flowers 2-whorled, rarely solitary; pedicels 8-15 mm long, angular, striate, thickened upwards, scarcely pubescent to ± glabrous, ± erect or curved; in fruit elongated to 30 mm and slender. Calyx 2.5-3 mm long, shortly cupular, 5-ribbed, 5-dentate, subglabrous; teeth 0.5-0.8 mm long, apically thickened and somewhat obtuse to ± subulate, spreading; in fruit enlarged and surrounding the base of it. Corolla greenish-white to dirty-white, rotate-campanulate; limb 8-9 mm across; lobes 1.5->2.5 mm long, ovate-oblong or ± triangular, obtuse or slightly acuminate, ciliolate. Filaments 1-1.5 mm long; anthers yellow, 1.7-1.9 mm long, oblong. Ovary c. 1 mm long, ± ovoid, rounded distally, glabrous; style 3.5 mm long, slightly thickened into a small stigma. Fruit scarlet, erect, glossy, (7)9-10 × (5)6-7 mm, globose-ovoid or broadly ellipsoid, rounded distally, smooth, glabrous. Seeds pale brownish, 3.5-4 × 2.5-3 mm, ovate in outline or ± reniform. [taken from Flora Zambesiaca (2014) for wild species, C. baccatum var. baccatum].
The domesticated form, var. pendulum, of this lowland South American species has cream-coloured flowers with paired gold or green markings. Typically, fruit are elongate with cream-coloured seeds.

Distribution

The species C. baccatum originated from Peru and southern Bolivia with varying distributions; the domesticated form var. pendulum spans across tropical South America, while the wild var. baccatum is less widespread and ranges from Peru to Brazil (Eshbaugh, 1970; Basu and De, 2003). According to Russo (2012), the wild taxon is common in Bolivia and northern Argentina with outlier populations in Peru and Paraguay.
The species is included in the Vascular Plants of Ecuador (2014) which, however, notes its presence in the country is "questionably reported from the Galapagos Islands on the basis of a single collection" (Wiggins and Porter, 1971).

Distribution Map

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Distribution Table

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History of Introduction and Spread

Among the first domesticated plants of Mesoamerica, Capsicum has been known since the beginning of civilization in the Western hemisphere and has been part of the human diet since 7500 BC (Basu and De, 2003). It was either Christopher Columbus or his accompanying physician Chanca who first reported the use of Capsicum in the Americas to Europe around 1493-1494 and certainly Columbus who introduced it across the Atlantic; by mid-17th century Capsicum was being cultivated throughout southern and middle Europe as a spice and medicinal drug, with introductions of one species to Japan and five to India (for mass cultivation in the colonies, from the Portuguese) around this time (Basu and De, 2003). At the time of Cristopher Columbus’ discovery of the Americas, the cultivated form of C. baccatum var. pendulum was only found in areas east and west of the Andes (Russo, 2012). Today the cultivated form, C. baccatum var. pendulum, has worldwide distribution and is reportedly the most widely domesticated pepper of Peru, even over C. annuum (Russo, 2012). The cultivated species has also been described as the most consumed species in Brazil (Spiller et al., 2008).
In the West Indies, Capsicum had been introduced to Jamaica by 1871, as Macfadyen observed the use of Capsicum fruit by Caribbean people as a food and drink condiment, but the plant is not mentioned by species (Macfadyen, 1871). The species was reported by name in Puerto Rico in 1881, during which Bello observed many forms being cultivated for culinary uses (Bello Espinosa, 1881). The species was present in Bermuda in 1918, as Britton reported it to be an occasional native plant occurring in rocky woodlands, and occasionally in gardens (Britton, 1918). The species can now be found across the Cayman Islands, Hispaniola, Jamaica, Puerto Rico, and the Virgin Islands, and is now considered an invasive introduction to Cuba (Oviedo-Prieto et al., 2011; Acevedo-Rodriguez and Strong, 2012). 

Risk of Introduction

Risk of introduction of C. baccatum is currently low to moderate, but the species possesses both desirable and undesirable traits that could cause potential invasiveness if not monitored, and further research is needed. The species is listed as a weed in the Global Compendium of Weeds (Randall, 2012) and is reported to be invasive to Cuba (Oviedo-Prieto et al., 2011) and a weed in Trinidad and Brazil (Holm et al., 1979; Randall, 2012). Invasive traits include tolerance of a wide range of precipitation and soil types, the species’ production of seeds viable for more than one year, wide distribution outside of its native range, and its ability to tolerate both acidic and alkaline soils (Basu and De, 2003; Ravishankar et al., 2003). Considering that the species has been cultivated in South America since pre-Columbian times (Basu and De, 2003) and is now present across tropical and subtropical regions around the world, the risk of introduction for this species may rise and should be monitored, especially in places where the species is cultivated.

Means of Movement and Dispersal

C. baccatum spreads by seeds. Dispersal is primarily through intentional and accidental introduction, as the species continues to be cultivated around the world for human consumption as a food, spice, and medicinal ingredient (Basu and De, 2003); the domesticated species is reportedly the most widely domesticated pepper of Peru (Russo, 2012) and has been described as the most consumed species in Brazil (Spiller et al., 2008). In Oceania the species reportedly escaped cultivation but has been grown as both a spice and ornamental plant; in Vanuatu it is used first and foremost as an ornamental shrub and as a boundary marker, which may contribute to spread of the species through accidental introduction and cultivation escape (Walter et al., 2007).

Pathway Causes

Pathway Vectors

Habitat

C. baccatum is primarily cultivated as a food and spice crop in agricultural and garden settings. In Zambia, it grows in disturbed ground and old cultivations as well as on anthills (Flora Zambesiaca, 2014). In Bermuda the species has been reported to grow in rocky woodlands and occasionally in gardens (Britton, 1918). In Peru, the domesticated form C. baccatum var. pendulum reportedly occurs in disturbed areas of the Andes and Amazonian regions, at altitudes 0-1500 m (Peru Checklist, 2014).

Habitat List

CategorySub categoryHabitatPresenceStatus
Terrestrial    
TerrestrialTerrestrial – ManagedCultivated / agricultural landPresent, no further detailsProductive/non-natural
TerrestrialTerrestrial – ManagedManaged forests, plantations and orchardsPresent, no further detailsProductive/non-natural
TerrestrialTerrestrial – ManagedDisturbed areasPresent, no further detailsProductive/non-natural
TerrestrialTerrestrial ‑ Natural / Semi-naturalNatural forestsPresent, no further detailsProductive/non-natural
TerrestrialTerrestrial ‑ Natural / Semi-naturalRocky areas / lava flowsPresent, no further detailsProductive/non-natural

Biology and Ecology

Genetics

Sporophytic chromosome count for the species is 24 (IPCN Chromosome Reports, 2014).

Reproduction

The flowers are protogynous, but readily self pollinate. In the field, high rates of outcrossing (up to 90%) can occur with insect pollination. Capsicum exhibits no inbreeding depression. The stigma is positioned slightly below the level of the anthers or exserted slightly beyond, in which case the chances for cross-pollination are greater. Pepper breeders and seed producers use caution when producing a seed crop to prevent uncontrolled cross-pollination. The flowers are normally solitary in the axils of the branches, with the occasional cluster type that causes multiple flowers to form at a node. Many of the wild species have multiple flowers per node. There are two to four or more locules within the fruit. The locules are separated by the placentae where the capsaicinoids are produced. The outer wall, or pericarp, is fleshy and varies in thickness. It consists of a very thin cuticle, five to eight compact layers of small collenchyma cells that are cutinized during maturation and provide a tough, colourless, epidermal layer or skin. The growth of the fruit in the early stage consists of rapid cell multiplication; in the later stages growth is chiefly by enlargement of the cells already formed.

Environmental Requirements

Capsicum plants grow best at low-to mid elevations with 7-8°C, annual rainfall of 300-4600 mm and well-drained, sandy or silt-loam soil with pH of 4.3-8.7 (Basu and De, 2003; Ravishankar et al., 2003). FAO reports optimal annual rainfall levels for C. baccatum var. pendulum to be 600-1250 (absolute 500-1500) and temperature absolutes of 15-32°C, with the ability to grow in climatic zones ranging from tropical wet or wet and dry, to subtropical humid, dry summer, or dry winter (FAO EcoCrop, 2014). In Bolivia, C. baccatum grows in dry to montane forest and dry valleys at altitudes of 0-2000 m (Bolivia Checklist, 2014), while in Paraguay, the species occurs in low and high-altitude forests (Paraguay Checklist, 2014).

Climate

Climate typeDescriptionPreferred or toleratedRemarks
Af - Tropical rainforest climate> 60mm precipitation per monthPreferred 
Am - Tropical monsoon climateTropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))Preferred 
As - Tropical savanna climate with dry summer< 60mm precipitation driest month (in summer) and < (100 - [total annual precipitation{mm}/25])Preferred 
Aw - Tropical wet and dry savanna climate< 60mm precipitation driest month (in winter) and < (100 - [total annual precipitation{mm}/25])Preferred 
Cs - Warm temperate climate with dry summerWarm average temp. > 10°C, Cold average temp. > 0°C, dry summersPreferred 
Cw - Warm temperate climate with dry winterWarm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)Preferred 

Soil Tolerances

Soil texture > light
Soil texture > medium
Soil reaction > acid
Soil reaction > neutral
Soil reaction > alkaline
Soil drainage > free

Notes on Pests

Biotic pathogens that cause problems include bacteria, fungi, mycoplasmas, virus, insects and nematodes.
Some of the most common bacterial diseases are bacterial spot (Xanthomonas campestris pv. vesicatoria), bacterial canker (Corynebacterium michiganense), bacterial soft rot (Erwinia carotovora pv. carotovora) and bacterial wilt (Pseudomonas solanacearum).
Fungi are one of the largest groups of organisms that cause diseases on pepper. The most important in terms of economic damage worldwide are anthracnose (Colletotrichum spp.), Cercospora leaf spot (Cercospora capsici), damping-off/seedling disease, early blight (Alternaria solani), grey mould (Botrytis cinerea), Phytophthora (Phytophthora capsici) and Verticillium wilt (Verticillium dahliae), the latter primarily a problem in temperate climates. Powdery mildew (Leveillula taurica) occurs rarely in cool climates, but is prevalent in warm climates and greenhouses.
In the tropics, viruses are the most serious disease problem and close to 45 viruses have been reported to infect peppers. Of these, more than half are transmitted by aphids. The other viruses are transmitted by nematodes, thrips, leafhoppers, whiteflies, beetles, fungi or by the grower handling infected plants. Some of the most widespread viral diseases are Cucumber mosaic virus, Pepper mottle virus, Potato virus Y, Tobacco etch virus, Tobacco mosaic virus and Tomato spotted wilt virus.
The insect pests most common to pepper plants are cutworms, aphids, pepper weevils, maggots, flea beetles, hornworms and leafminers. The major insect pest species vary by geographic region. In the north-eastern and central USA and Canada, European corn borer (Ostrinia nubilalis) is the primary pest species.
Early in the season, cutworms are the most damaging pests to both seeded and transplanted peppers. Seeded peppers are also subject to attack by flea beetles when the cotyledons emerge. Green peach aphids (Myzus persicae) can become numerous at any time, but are probably more prevalent during the summer. Occasionally, loopers will feed on the foliage, exposing the pods to sunscald. In warmer climates, fall and beet armyworms (Spodoptera frugiperda, S. exigua), yellow-striped armyworms (S. ornithogalli), corn earworm (Helicoverpa zea) and variegated cutworms (Peridroma saucia) may feed on pods and cause the pods to drop or become unmarketable. The beet armyworm will feed on the foliage. Hornworms (Manduca spp.) and the cabbage looper (Trichoplusia ni) are generally considered minor pests because they feed on foliage and do not enter the fruit.
Spider mites (Tetranychus spp.), various nematodes and animals can also cause damage. Peppers are also subject to attack by various nematodes. Among these are species of Meloidogyne and the root-lesion nematode, Pratylenchus penetrans. Three species of root-knot nematodes cause serious damage to peppers, Meloidogyne incognita, Meloidogyne hapla and Meloidogyne arenaria.
It is common in rural areas that herbivores such as deer, rabbits and mice destroy peppers. Birds can be pests by pecking holes in pods to get at the seeds.
Weed control has to start as pre-emergence treatment by cultivating or by using herbicides. After establishment, weeding can be done mechanically, although in the tropics it is often manually. An effective method of weed control is covering the soil with organic or plastic mulch.

List of Pests

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Non-Infectious Disorders

Physiological disorders

The two most common abiotic disorders are blossom-end rot and sunscald. Blossom-end rot occurs when the plant is unable to translocate adequate calcium to the fruit, a condition caused by fluctuating soil moisture (drought or over watering), high nitrogen fertilization or root pruning during cultivation. This disorder first appears as a water-soaked area on the fruit. The tissue near the blossom end of the pods has a brown discoloration. Unlike tomatoes, blossom-end rot is never actually at the blossom end, but more on the side of the fruit. Sunscald is caused by intense sunlight on fruit that had been growing in the plant’s shaded canopy. The smaller-podded varieties with erect fruit are not as susceptible to sunscald as the large-podded varieties, such as bells and New Mexican pod types.
Another common disorder is flower drop. Flower buds, open flowers and immature pod drop are caused by a variety of conditions. Heat stress, insufficient water and excessive or deficient nutrient levels have been reported as casual agents.
Stip or blackspot is a newly discovered abiotic disorder on pepper fruit. Stip causes grey-brown to greenish spots on the fruit, most noticeable on red fruit that matures in the autumn. The disorder manifests itself when peppers are grown under cooler temperatures and associated with a suspected calcium imbalance, and possible shorter day-length. Stip can occur in the interior tissue of fruit as well as on the external surface. The disorder only affects some pepper cultivars, so the best control is to plant resistant cultivars.
Oedema appears as numerous small bumps on the lower side of the leaves, and sometimes on the petioles. The cause is most likely to be over-watering, although high humidity can also contribute to the problem. Control measures include reduced watering and better air circulation around the plant.
Inadequate phosphorus produces weak plants with narrow, glossy, greyish-green leaves. The red or purple colouration of stems and leaves often associated with phosphorus deficiency does not develop on peppers. Fruit produced on pepper plants deficient in phosphorus are shorter. Low levels of potassium produce a bronzing condition on the pepper leaves, followed by necrosis and leaf drop. Magnesium deficiency is characterized by pale-green leaf colour followed by interveinal yellowing, leaf drop, small plants and undersized fruit. Magnesium deficiency generally occurs when pepper plants are grown on acidic, sandy soils in areas of high rainfall. Compared with soil applications, spray applications of magnesium and other minor elements are more effective and have a more rapid (but shorter-lived) effect.

Impact Summary

CategoryImpact
Economic/livelihoodPositive and negative

Impact: Environmental

C. baccatum is classified as a weed in the Global Compendium of Weeds (Randall, 2012), indicating its potential threat to the environment. Little data is currently available on the impacts of C. baccatum becoming weedy or invasive, and this should be researched in order to prevent and control the problem proactively. 

Risk and Impact Factors

Invasiveness

Proved invasive outside its native range
Abundant in its native range
Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
Pioneering in disturbed areas
Tolerant of shade
Has propagules that can remain viable for more than one year

Likelihood of entry/control

Highly likely to be transported internationally deliberately
Difficult to identify/detect in the field

Uses

C. baccatum has been valued by cultures around the world for its culinary use as a spice and vegetable, as well as in medicine and as an ornamental. In Oceania the species reportedly escaped cultivation and has been grown as both a spice and ornamental plant; in Vanuatu it is used first and foremost as an ornamental shrub and as a boundary marker (Walter et al., 2007). C. baccatum is also used as the ‘pepper’ chemical ingredient in self-defense pepper sprays (Basu and De, 2003). Pharmacological studies have also shown the species possesses anti-inflammatory properties and potential for pharmaceutical development (Spiller et al., 2008).

Uses List

Environmental > Boundary, barrier or support
Materials > Chemicals
Medicinal, pharmaceutical > Source of medicine/pharmaceutical
Medicinal, pharmaceutical > Traditional/folklore
Human food and beverage > Food additive
Human food and beverage > Spices and culinary herbs

Cultivation

Transplanting of seedlings into the field takes place about 35 days after sowing, when the plants have reached the eight-true-leaves stage. The young plants are transferred to their final positions when they are approximately 10-12 cm high. They are usually planted out in rows 60-80 cm apart and 35-45 cm apart within the rows, depending on the vigour of the cultivar. Where furrow irrigation is planned, the ridges are prepared during site preparation. In areas where the climatic conditions are ideal, growers sow direct at the same inter- and intra-row spacing and in this method three or four seeds are sown at each station. The emerged seedlings are subsequently thinned to one strong seedling per station. Many cultivars require staking to avoid lodging. Flowering begins 2-3 months after planting.

Site preparation and planting

Ideally, peppers should not be planted in the same field more than once every 3-4 years and, in the intervening years, the crops grown in the field should be non-solanaceous crops such as wheat, brassicas, maize, lucerne and legumes. The use of raised beds is a common cultural practice for pepper production, but it is not always essential. Factors such as soil type, soil salinity, cultivation methods, type of irrigation, and drainage need to be considered when deciding on whether or not raised beds will be used. Most commercial pepper production in the USA occurs on 15-20 cm raised beds to promote drainage. Beds can be various sizes but are commonly 122 cm wide and 183 cm centre-to-centre. Most peppers are grown on soil that is highly prepared with tillage work. The soil can be tilled in a number of ways and rotovators can create a smooth bed if there is little residue. In some cases, fields are ploughed, particularly if heavy residue must be incorporated, tilled, and the beds reshaped. Cultural factors are implicated as being more important in seedling survival rate than the physical, mechanical transplanting operation. The optimum soil pH is 6.0-6.5, if found to be lower than this a suitable liming material should be applied during site preparation at a rate of 30 g/m2.

Pruning, training and thinning

Improved methods of transplant production and handling have increased survival rates of relatively young transplants, and field pruning to control growth of large pepper transplants is seldom used. Staking of field grown plants has been used to improve yield but its effectiveness depends on N and water supply.
Young transplants naturally branch into two or occasionally three shoots. This occurs after the fifth to the eighth node. Under greenhouse conditions, plants are normally trained to two stems each. Peppers are brittle so they need adequate support, the most common support system being string. Strings are tied to the stems and the string is supported from overhead wires 2.5-3.0 m above. At about 4 weeks after planting, two of the strongest shoots are selected. Stems are tied to the strings supported from the overhead wires. The stem requires twisting around the string stock wire every 10-14 days. Failure to maintain regular pruning can result in lost production and reduced plant growth rates. Ideally, a pepper plant will set one fruit for every two leaves. When the lateral branches have four leaf axils above the first fork, the flowers are allowed to set. Misshapen and diseased fruit are removed as soon as possible. Side shoots are removed as soon as possible, allowing for better light penetration and larger flower development. The option of leaving more leaves per shoot is possible when light intensity is high, the additional leaves will prevent sunscald of the fruit. The first flower on the young pepper plant is removed. The removal is important because if a fruit sets, early extension growth is reduced and the plant will have a stunted habit with a poorly developed fruit.

Irrigation

Pepper plants have extensive root systems that are moderately deep-rooted with taproots reaching a depth of more than 1 m. Although peppers are generally drought resistant, intermittent periods of moisture, and/or nutritional stress can dramatically reduce plant growth, fruit size, and yield. Flowers and young fruit may abscise if water stress occurs during flowering. The water requirements for pepper production range from 400 to 1000 mm. The broad range reflects differences in cultivar and environmental conditions. The general irrigation recommendation of a minimum of 2.5 cm/week applies to pepper. On sandy soils, more water may be required. Drip irrigation provides uniform and efficient water application directly to the root zone. Furrow irrigation may be an option in areas where water is plentiful and the infrastructure supports it.

Plant nutrition and fertilizers

Peppers do not remove as many nutrients from the soil as tomatoes. When both the plants and fruit are considered, a pepper crop will remove approximately 134 kg N, 13.4 kg P, 134 kg K/ha. Fertilization should be performed in response to both soil and foliar testing to ensure that only the nutrients needed by the plant are applied. Peppers respond to N fertilization, which is often applied prior to transplanting by row banding. Depending on testing results, a second application of N may be applied again at first flowering. However, excessive N wastes costly fertilizer, may damage the environment, and favours vegetative growth while inhibiting reproductive growth, thus delaying maturity and decreasing marketable yields.
Fertigation in the field is common with plasticulture pepper production. Phosphorus fertilizer may be banded at transplanting because of limited solubility and the rest of the nutrients fertigated during the season in response to tissue or petiole sap mineral analysis. A liquid starter solution high in P is often applied to the root zone at transplanting to stimulate early growth. A standard starter solution recipe may consist of 1.5 kg/200 litre of 8N-24P-8K or 10N-52P-17K, with 118 ml applied to each transplant.

Soil management

Field grown peppers are often produced with some type of mulch and the most common artificial mulching material is plastic (usually polyethylene) film. Black plastic mulch typically improves pepper yield compared to bare soil although negative effects may occur under high ambient temperatures. The main advantages are increased early yields, improved moisture retention, inhibition of weeds, reduced fertilizer leaching, decreased soil compaction, fruit protection from soil deposits and soil microorganisms, and facilitation of fumigation.
Weed control has to start as pre-emergence treatment by cultivating or by using herbicides. After establishment, weeding can be done mechanically, although in the tropics it is often done manually.

Protected cultivation

Peppers production in glasshouses and other protective structures is significant primarily in areas where crops cannot be matured outdoors. Indeterminate cultivars developed for greenhouse production are carefully trained vertically on strings and pruned to efficiently use greenhouse space. The Dutch greenhouse system uses fertigated rockwool media similar to the system used for greenhouse tomato production. Night temperatures should be maintained at <15°C for best fruit development. Bees or mechanical agitation are used to improve greenhouse pollination. A bumble bee hive containing approximately 60 bees per 1440 m2 provides sufficient greenhouse pollination. Manual pollination by workers is generally less efficient and considerably more expensive.

Harvesting

Harvesting

The majority of peppers in the world, destined for fresh market, are harvested by hand. The reason for hand harvesting is quality. Hand-picked peppers are of a higher quality because humans can instantaneously reject mouldy, underripe, overripe or damaged pods. There is no abscission zone, so the pedicle of fruits must be carefully cut or expertly snapped by hand without damaging the brittle branches and surrounding leaves. Humans also do not pick as many leaves and stems as machines and there is an increased yield per unit area. The increased yield is associated with less damage to the plants as human pickers move through the field. Machine harvesters cause more damage and machine harvested plants take longer to recover and set more fruit. Mechanical harvesting is more common for processing crops because fruit damage is less critical if no storage is needed before processing. Peppers for processing are destructively harvested either by hand or machine. For small scale production of chilli powder, paprika, and other dried pepper products, entire plants with fully coloured fruit are cut and field dried when environmental conditions are favourable.

Harvesting date

Picking the fruits may start 3-4 months after planting. The harvest time depends on the desired product: fruits are sometimes picked when they are unripe and green, and others are picked ripe, when the fruits are red, yellow, orange or brown. The harvesting of peppers is flexible and based on colour, fruit size, and/or wall thickness. Generally, the surface of older fruit is firmer, shinier, and waxier than younger fruit. More mature fruit tend to resist shrinkage after harvest, more than less-mature fruit whose cuticle is not fully developed. Green fruit are harvested after reaching their mature size, but before physiological maturity and colour change occur. Fruit are physiologically mature when the seed they contain are fully developed and they obtain their mature colour, which is often red or yellow. Some growers use accumulated heat units for scheduling harvests.

Mechanized harvesting

For large-scale production, mechanical harvest of the red chilli and paprika crops has been widely adopted in recent years, with more than 80% of the commercial crop in the USA harvested by machine. There are several different types of mechanical harvesters currently used for red chilli and paprika. The three most common picking heads are the finger-type, the belt-type, and the double helix. The finger-type head is designed with a series of counter- rotating bars with fingers. The fingers comb the plants to strip off the pods on to conveyor belts. The mechanism is aggressive and may harvest unwanted plant debris. A belt-type harvester has two sets of counter-rotating vertical belts imbedded with fingers that comb both sides of a plant. The most widely used picking head is the double-helix design. The helices may be vertical or oriented at an angle. The helices rotate in opposite directions to each other, snapping pods off of the plants and flipping them on to conveyor belts on either side.

Yield

Under greenhouse production, well-grown and managed crops of Capsicum on average can produce yields of approximately 15 kg/m2. There is no information, however, available on yields of field grown C. baccatum. However, FAOSTAT (2016) reported yields of green chillies and peppers for South American countries where is it is the most widely grown Capsicum spp., may give an indication of its yield. In 2013, average yields were highest in Chile (60.98 t/ha), followed by Peru (9.28 t/ha), Ecuador (3.17 t/ha) and Bolivia (2.82 t/ha).

Postharvest Treatment

Postharvest treatment and handling

Fresh market Fruits are susceptible to mechanical injury and may be easily damaged during shipment. After harvest, fruit for fresh market should be rapidly cooled to about 10°C to remove field heat. Peppers are washed in ambient or warm chlorinated (300 ppm) water to reduce postharvest diseases before storage or shipment.
Dried Peppers harvested for drying are dried using sun drying in the field or larger, commercial operations employ either tunnel or belt dryers. Both systems are more reliable and sanitary than sun drying. However, these methods use more energy, especially early in the harvest season when succulent pods are harvested and must be dried. Another drawback to tunnel and belt dryers is their limited capacity. The dehydrated pepper fruit is ground and rehydrated to 8-11% moisture, an optimal level for storage. Antioxidants such as ethoxyquin are sometimes added to reduce colour loss. Cold storage at 3°C is recommended.
Canning The most common pod types to be canned are New Mexican, pimiento and jalapeno. Pepper fruit are normally processed at 100°C, but long exposure time to this thermal treatment or to pressure processing can excessively soften the fruit. Acidification of the product reduces the pH below 4.6, decreases the thermal resistance of microorganisms, allows the canner to reduce the thermal exposure time and process peppers at atmospheric pressures. The pH of peppers has been shown to vary among early to late season harvests, along with the degree of ripeness. Therefore, concentrations of acidulent must be adjusted to account for the pH variability of the raw fruit. Properly canned fruit generally have a maximum shelf life of 2 years. In addition to pH considerations, fruit texture is a major concern of processors. Fruit softening of canned peppers can be minimized with calcium treatments. Calcium chloride is the preferred calcium source for both pimientos and jalapenos, although calcium lactate is also acceptable. In canned pimientos, the combination of citric acid and 0.02% calcium chloride to the product significantly increased both firmness and drained weight when compared to either treatment alone. The firmness of canned jalapenos is increased, without imparting bitterness, when 0.2% calcium chloride is added to the product.
Brining Pepper fruit that are usually brined include cherry, wax, pepperoncini, jalapeno and serrano. The pickling or brining process involves adding sufficient amounts of salt and acetic acid to prevent microbial spoilage; peppers packed in this manner have superior textural qualities as compared to canned peppers. The effectiveness of brining for preservation is related to the rate of acid diffusion into all parts of the fruit and the time required to reach an equilibrium pH of 4.6 or below. The primary areas for acid penetration are through the stems and calyxes and into the placentas. The interior of fruit walls is the last area to become acidified, and the entire process can take a minimum of 6 days. Blanching can also improve the rate of acid penetration into the fruit and therefore reduce the pH variability among fruit parts.
In most commercial operations, peppers are brined in a two-step process. First, fresh pepper fruit are placed in a primary brine to firm and preserve the fruit. After a minimum period of 2-8 weeks, depending on cultivar and process, the fruit are removed from the first brine, washed, graded a second time and then re-packed in finishing brine. Only the best quality fruit are packed whole or sliced in the finishing brine. Vinegar and salt levels may be reduced in the finishing brine and spices may be added for flavouring. This second brine is usually added to the fruit in the final container, and the container is sealed and hot-packed. A large portion of fruit are never packed in the second brine, instead they are chopped and blended into other food products after the initial brining.
Preservatives are often used in the initial brine to further prevent fruit softening and discoloration. Sodium bisulfite (0.5-1% by weight) is the most common preservative in pickled pepper. Sodium bisulfite is an effective preservative, but it imparts an off-flavour that is leached in the second packing brine. Bisulfite has also been implicated in certain food allergies experienced by asthmatics. Calcium chloride (0.25-0.50% by weight) can replace bisulfites, but is less effective at firming the pods, can impart bitterness at high concentrations, darken fruit colour and requires additional salt for optimal rigidity of the fruit. Brined pepper fruit can be produced without preservatives by using a more costly refrigerated process. Cold temperatures allow the fruit to imbibe brine completely, while slowing the growth of bacteria that can soften the fruit. The first 4-8 weeks are the most critical period of the brining operation. If the conditions are not optimal, the fruit can soften. After 6-8 weeks in cold storage, the fruit reach an osmotic equilibrium with the brine solution and further softening does not occur. Brined fruit without chemical preservatives are then packed in finishing brines with higher vinegar and salt concentrations than those with bisulfite.

Storage

Fresh chilli peppers are not as chilling sensitive as bell peppers. Chilli peppers stored above 7.5°C suffer more water loss, shrivel, colour change, and decay. Storage at 7.5°C is considered the best for maximum shelf-life of 3-5 weeks. Chillies can be stored at 5°C for at least 2 weeks without visible signs of injury. Storage at this temperature reduces water loss and shrivelling, but after 2-3 weeks, chilling injury is mostly detected as discolouration of the seeds. Ripe or coloured chillies are less chilling sensitive than mature green chillies. Unlike tomatoes, peppers are not climacteric. Many chilli peppers do not ripen after harvest when treated with ethylene. Bell pepper ethylene production rates are very low in the range of 0.1-0.2 ml/kg/h at 10-20°C. However, some chillies, such as habaneros, show increased ethylene production during ripening and may produce over 1 ml/kg/h at 20-25°C. Responses to added ethylene depend on the particular type of chilli. Chilli ‘Poblanos’ for example may respond to ethylene treatment, while ‘Jalapeno’ peppers do not. As with bell peppers, holding partially coloured chilli peppers at warmer temperatures of 20-25°C with high humidity (>95%) is effective to complete colour development. Adding ethylene may further enhance ripening but the response is cultivar dependent. Red and yellow physiologically mature bell peppers are more expensive to produce than green because fruits must remain on the plant longer to develop their mature colour. This reduces total yields because mature peppers inhibit the set and growth of new fruit. Fruit are also susceptible to disease and insect attack during their extra time on the plant. Mature red fruit also have 10 times more provitamin A content, softer texture, and higher soluble sugar content, which also adds to their value. The red colour develops because of increased carotenoid synthesis and a breakdown of chlorophyll. As green bell peppers begin to turn to red, their colour first changes to brown during the transition because both the red carotenoids and green chlorophyll are in the fruit wall simultaneously. This is often called the “chocolate” stage. Once harvested, chocolate peppers will remain brown because they do not easily ripen after harvest. In some markets, chocolate peppers have less value because the colour is unappealing to consumers. High relative humidity is necessary to limit desiccation. Water-soluble wax is sometimes used to coat peppers to decrease shrivelling and increase storage life. However, waxing may increase the possible incidence of bacterial soft rot. Bell peppers are also wrapped with a thin layer of plastic film individually or grouped in a plastic tray to preserve freshness and reduce mechanical damage from handling. Waxed or film-wrapped peppers can be stored from 10-14 days at 13°C. Shelf life varies among cultivars. Deterioration is often due to moisture loss and some fruit types are more prone to desiccation than others. Peppers generally do not respond well to controlled atmosphere storage. Low O2 atmospheres (2-5% O2) alone have little effect on quality and high CO2 atmospheres (>5%) can cause pitting, discoloration, and softening, especially at temperatures >10°C. Atmospheres of 3% O2 + 5% CO2 are more beneficial for red than green peppers stored at 5-10°C.

Genetic Resources and Breeding

Genetics

Sporophytic chromosome count for the species is 24 (IPCN Chromosome Reports, 2014). C. baccatum has a longer karyotype length (and increased DNA content), 3-4 chromosomes with active nucleolus organizer regions, increased GC-enriched heterochromatin (7-9%), and a highly complex heterochromatin banding pattern compared to other Capsicum species (Moscone et al., 2007). A list of known genes can be very useful to pepper breeders, especially if a collection of germplasm that contains representative specimens is available. In 1965, Lippert and colleagues produced a gene list for pepper (Lippert et al., 1965) which included 50 genes and a standardization of rules for naming and symbolizing. Wang and Bosland (2006) provided the most recent update, adding 92 previously unreported genes to bring the total to 292.
The US National Plant Germplasm System houses an extensive Capsicum germplasm collection at the Southern Plant Introduction Station located in Griffin, Georgia. This gene bank has a worldwide collection of approximately 5000 Capsicum accessions. Globally, there are several other gene bank collections. Many of the accessions in each collection are duplicates of material maintained in other collections. The most active collections are the Asian Vegetable Research and Development Center (AVRDC) in Tainan, Taiwan, the Centro Agronomico Tropical de Investigations y Ensenanza (CATIE) in Turrialba, Costa Rica, the Centre for Genetic Resources (CGN) in Wageningen, the Netherlands, and the Central Institute for Genetics and Germplasm in Gatersleben, Germany. There are important South American collections of C. baccatum holding wild and domesticated forms. The collection of the Instituto Nacional de Innovación Agraria (INIA), Peru, holds 712 Capsicum accessions of which 70 are domesticated C. baccatum. The collection of the Centro de Investigaciones Fitoecogenéticas de Pairumani (CIFP) in Bolivia holds 487 Capsicum accessions of which 217 accessions are of domesticated C. baccatum and 11 are of the wild species C. baccatum var. baccatum. This latter collection conserves an important amount of C. baccatum genetic diversity compared to international Capsicum collections as Bolivia is a primary centre of diversity of cultivated C. baccatum var. pendulum (Zonneveld et al., 2015). Embrapa Clima Temperado, a research institution in southern Brazil, maintains a collection of 347 Capsicum accessions, including 286 of C. baccatum (Barbieri et al., 2007).
Throughout Latin America, it is not uncommon to find wild and semi-wild Capsicum species growing in close proximity to cultivated species, in domestic gardens or at the edges of cultivated fields. This situation allows introgressive hybridization to takes place between the domesticated species and closely related wild species or sub-species, spontaneously giving rise to novel or intermediate forms which are then selected and propagated by observant farmers in an ongoing process of on-farm crop evolution (Zonneveld et al., 2015).

Breeding objectives

These focus on fruit attributes including yield, colour, size, shape, and pungency, and fruit and foliar disease resistance in open field and protected culture. Varying levels of resistance to powdery mildew caused by Leveillula taurica have been reported in C. baccatum var. microcarpum and C. baccatum var. pendulum. C. baccatum is also a valuable species in Capsicum breeding, in particular as a source of disease resistance against anthracnose and powdery mildew.

Breeding methods

Capsicum breeders are faced with the arduous task of assembling into a cultivar the superior genetic elements necessary for increased yield, protection against production hazards and improved quality. Each type has its own unique set of characteristics that must be met in order to be commercially acceptable. Breeders are therefore faced with a plethora of unique objectives, and in order to achieve those objectives, they may use quite a few different breeding methods. The methods used are determined by the breeder to best fit the goals of the breeding programme.
The first pepper breeders, the indigenous people of tropical America, used mass selection to domesticate the five different species of Capsicum. Another breeding method used by many pepper breeders is the pedigree method, which involves keeping records of the matings and their progeny. A good cultivar can be made better by using the backcross method, which uses a successful cultivar as a recurrent parent following an initial cross between that successful cultivar and a separate individual that serves as a donor parent for a given desired character. Following successive backcrosses to the recurrent parent, the breeder arrives at a cultivar identical to the recurrent parent, but containing the additional desired trait from the donor parent. Several other breeding methods have been successful. For example, recurrent selection, where individual plants are selected from a population followed by intercrossing to form a new population. Mutation breeding is a means by which mutations are generated in peppers to improve economically important traits or to eliminate deleterious traits. It is not a major breeding method, but may be a means of producing novel mutants of interest.
Biotechnology is being used to develop new cultivars. For many years, Capsicum spp. were recalcitrant to the methods of genetic transformation. Reports that pepper has been successfully transformed using Agrobacterium tumefaciens has given impetus for commercial seed companies to begin programmes introducing novel genes into peppers. Since the late 1980s, efforts to tag and map identified genes using molecular methods have provided markers linked with important traits for use in marker-assisted breeding programmes for sweet and hot pepper. Utilizing mapping populations developed from a series of interspecific crosses, RFLP, AFLP, RAPD, and SSR markers have been integrated to improve the average marker density and facilitate identification of simply inherited and complex attributes.

Major Cultivars

‘Aji Amarillo’, sometimes called 'Yellow Pepper', 'Yellow Peruvian Pepper' or ‘Tscabeche' in the USA, is the most common C. baccatum pod type in Peru. Its pods are 10-15 cm long and a deep orange when mature. They are thin-fleshed and have a fruity flavour with berry overtones and a searing, clear heat. ‘Aji Ayucllo' is a wild pepper variety found in the Peruvian rainforest, near the Chanchamayo and Villa Rica Valleys. The fruits are small, thick fleshed and oval, with a moderate heat. ‘Aji Norteno' or the 'northern aji' is popular in the northern coastal valleys of Peru, with production centred on the Vird and Lambayeque valleys, about 1000 km north of Lima. The fruits, which mature to yellow, red or orange, are 8-10 cm long and 2 cm wide. They are slightly curved and taper to a point.
Other cultivars and varieties of C. baccatum include ‘Brazilian Starfish’, ‘Peppadew’, ‘Bishop’s Crown’, and ‘Lemon Drop’ (World of Chillies, 2016). In addition, ‘Aji Ethiopian Fire’, ‘Aji Fruto’, ‘Fatalii Gourmet Ajo Fantasy’, and ‘Sugar Rush’ are available (UK Chilli Seeds, 2016).
Principal sources: Bosland and Votava (2012)

Propagation

Capsicum plants are propagated from seeds, as asexual propagation is not used in commercial production. Both open-pollinated and F1 hybrid cultivars are commonly used. The crop can be established in the field by direct seeding, by containerized transplants grown in multicellular trays in the greenhouse, or by using bare-root transplants that are field grown. Each method has advantages, and each is suitable for specific production systems. Direct seeding requires less labour and is less costly. However, with new hybrid cultivars costing 10 to 20 times more than the cost of open-pollinated cultivars, transplanting to a field stand is the only feasible option. However, direct seeding is still used for large-scale production of mechanically harvested peppers for processing as it makes for higher density plantings. High quality seeds germinate in 6-10 days at 15.6-29°C. C. baccatum non-dormant seeds also germinate fully at 10 or 13°C. The presence or absence of light is not a factor in Capsicum seed germination. Seed enhancement technologies such as seed coating (pelleting and film coating), controlled hydration (priming) or gel coatings containing growth-promoting substances such as gibberellic acid can improve the speed and synchrony of pepper seedling emergence. Seed priming in particular can effectively reduce the mean time to germination, particularly at low temperatures, and is a commonly used commercial seed treatment.

Nutritional Value

Capsicum fruit vary in size, shape, colour, flavour and pungency, and this variation is also reflected in their nutritional composition. The nutritional composition is determined by the species, the cultivar, the growing conditions and fruit maturity. Further changes can occur during postharvest handling and storage, but peppers are extremely good sources of many essential nutrients. Under both greenhouse and open field conditions in Spain, C. baccatum demonstrated considerable variation in composition and level of red and yellow carotenoids (CR and CY, respectively), ascorbic acid, and total phenolics. These are important antioxidants with benefit for human health. Several accessions had similar or higher levels than C. annuum indicating that C. baccatum is a valuable source of antioxidant compounds (Rodríguez-Burruezo et al., 2009).
According to the USDA National Nutrient ​Database, raw red hot chilli peppers contain (per 100 g fresh weight), approximate values of 1.87 g protein, 0.44 g total lipid (fat), 8.81 g carbohydrate (by difference), 1.5 g fibre (total dietary), 5.30 g total sugars, 88.02 g water, and 40 kcal. In addition they are rich sources of vitamin C (143.7 mg), vitamin A (48 µg), calcium (14 mg), iron (1.03 mg), thiamin (0.072 mg), riboflavin (0.086 mg), and niacin (1.244 mg). Peppers also contain the provitamins α-, β- and γ-carotene and cryptoxanthin, all of which are transformed in the human liver into vitamin A.
The lipophilic ingredient responsible for the burning sensation and pungency of chillies, capsaicin (methyl vanilly nonenamide), is present in varying amounts in different types of peppers and genotypes within species. The fruit of wild species along with most pod types are pungent. The production of capsaicinoids is controlled by a single recessive gene. The Scoville Heat Unit (SHU) is an index used to measure pungency. Peppers are classified into five main groups based on pungency level: non-pungent or paprika (0-700 SHU); mildly pungent (700-3000 SHU); moderately pungent (3000-25,000 SHU); highly pungent (25,000-70,000 SHU); and very highly pungent (>80,000 SHU).
Since pre-Columbian times, Capsicum spp. have been used as medicinal plants. Today, peppers are one of the most widely used of all natural remedies. It may have been the plants' medicinal uses that caused the indigenous peoples of the Americas to domesticate peppers. The Mayas and Aztecs used Capsicum to treat throat and lung ailments and to relieve toothaches.
Capsaicin is currently being used to alleviate pain, and is the most recommended topical medication for arthritis. At nerve endings, a neurotransmitter called substance P informs the brain that something painful is occurring. Capsaicin causes an increase in the amount of substance P released. Eventually, the substance P is depleted, further releases from the nerve endings are reduced, and the pain experienced by the patient is decreased. A decrease in substance P also helps to reduce the long-term inflammation that can cause cartilage to break down. A cream containing capsaicin is also used to reduce post-operative pain in mastectomy patients, and to reduce 'phantom limb' pain in amputees. Prolonged use of such cream has also been found to help reduce itching in dialysis patients, the pain from shingles (Herpes zoster), and cluster headache.
Spiller et al. (2008) demonstrated antiinflammatory effects of the juice of red fruits of C. baccatum on carrageenan- and antigen-induced inflammation in rats. They suggest that the antiinflammatory effect may be due to the inhibition by capsaicin of pro-inflammatory cytokine production at the site of inflammation.

Production and Trade

According to FAOSTAT (2016), world production of chillies and peppers (Capsicum spp.) in 2013 was 31,116,944 t (fresh) and 3,446,634 t (dried). The leading producers of fresh chillies and pepper were China (15,823,000 t), followed by Mexico (2,294,400 t), Turkey (2,159,348 t), Indonesia (1,726,382 t), Spain (999,600 t) and the USA (889,269 t). India produced nearly 40% of the world production of dried chillies (1,376,000 t), followed by China (300,000 t) and Peru (164,000 t). India exported 290,448 t, 21% of its dried chilli production.
Leading exporters of fresh chillies and peppers in 2013 were Mexico (793,501 t), Spain (583,827 t) and the Netherlands (407,823 t). The Netherlands is one of the world leaders in intensive greenhouse pepper production. The United States was the leading importer (905,822 t) followed by Germany (359,627 t).
World production figures for C. baccatum are unavailable; however, it is the domesticated pepper of choice in Bolivia, Ecuador, Peru and Chile, and the most commonly grown Capsicum species in South America. FAOSTAT (2016) fresh Capsicum spp. production and area harvested figures for 2013 for these South American countries are: Bolivia (15,448 t and 5,461 ha), Ecuador (5,704 t and 1,796 ha), Peru (13,000 t and 1,400 ha) and Chile (39,000 t and 609,851 ha).

Prospects

The study of pepper metabolites is a rapidly expanding field, a result of pepper's importance as a vegetable crop and staple ingredient in many cuisines. Capsicum fruit contain a rich diversity of metabolites that are reported to have antioxidant, hypoglycaemic, immunogenic, antihypertensive, anticholesterol, antiinflammatory, and antimutagenic effects. Given the wide variation for individual compounds in different species and genotypes, pepper lends itself to the enhancement of its qualities through manipulation of metabolite levels using conventional breeding and nonconventional approaches.
Future prospects for application of biotechnological tools are promising in Capsicum, especially for new hybrids developed by tissue culture and biotechnology methods. There has been a growing expectation that the biotechnology industry will deliver a second generation of transgenic products for more challenging traits relating to yield and yield stability, which are under complex polygenic control.

Gaps in Knowledge/Research Needs

Less is known about this species compared with other members of the Capsicum genus, and further research is recommended in the areas of the potential environmental, social and economic impact if the species becomes invasive. As it is only reported to be invasive to Cuba and Trinidad, additional data on the species’ invasive or weed status would help to steer future actions for monitoring and, if necessary, control.

Links to Websites

NameURLComment
Catalogue of Seed Plants of the West Indieshttp://botany.si.edu/antilles/WestIndies/catalog.htm 
GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gatewayhttps://doi.org/10.5061/dryad.m93f6Data source for updated system data added to species habitat list.

References

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Published online: 4 December 2014

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Marianne Jennifer Datiles
Pedro Acevedo-Rodríguez

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