The Capsicum Crop: An Introduction

1
Pasquale Tripodi and Sanjeet Kumar

Abstract we provide an overview of the origin

Capsicum (Capsicum spp.), also called as and history of the pepper, describing its
pepper, is a main vegetable and spice crop economic importance, properties, and
originated in the American tropics and commercial mar- ket types.
today cultivated all over the world for
fresh, dried, and processing products.
Around the genus
Capsicum there is an increasing interest and
fascination due to the considerable 1.1 Origin and Diffusion
variation for several traits, which makes
this crop extremely versatile and suitable The genus Capsicum is part of the large Sola-
for innumer- able uses as food and non-food naceae family, which, among the more than
products. The genus Capsicum includes over 90 genera and 2500 species of flowering
30 species, five of which (C. annuum, C. plants, includes commercially important
frutescens, C. chi- nense, C. baccatum, and vegetables such as tomato, potato, and
C. pubescens) are domesticated and mainly eggplant. This genus is native to tropical and
grown for consump- tion. A large number of subtropical America (Hunziker 2001) in a
accessions of domes- ticated and wild wide region comprising Mexico and northern
species are stored in the world seed banks, Central America, the Car- ibbean, the lowland
representing a valuable resource for Bolivia, the northern low- land Amazonia, and
breeding in order to transfer traits related to the mid-elevation southern Andes, where
resistances to various abiotic and biotic archaeological evidence suggests use of this
stresses as well for quality improvement. spice crop since 6000 BC (Davenport
The recent advances in terms of genetic 1970; Basu and De 2003; Perry et al. 2007).
and genomic knowledge will help to At the beginning, fruits were exchanged for
unlock the potentiality of these resources. In black pepper (Piper nigrum), a species similar
this chapter, in taste (though not in appearance) although not
phylo- genetically related to Capsicum
(Gordo et al.
P. Tripodi (&)
2012). For this reason, it was incorrectly
CREA Research Centre for Vegetable and named
Ornamental Crops, 84098 Pontecagnano Faiano, “pepper” (Walsh and Hoot
Italy 2001).
e-mail: [Link]@[Link]
It was Fuchs, who proposed for the first
S. Kumar time in 1543, the botanical term Capsicum, which
World Vegetable Center, Post Box 42, Shanhua,
Tainan 74199, Taiwan was adopted later in 1753 by Linneo. The
name would be the Neolithic derivation of
Greek “Capsa,” which refers to the peculiar
shape of the
© Springer Nature Switzerland AG 2019 1
N. Ramchiary and C. Kole (eds.), The Capsicum Genome, Compendium of Plant Genomes,
[Link]
 21 The Capsicum Crop: An P. Tripodi and S. 2
fruit. The crop was firstly introduced in Europe
types and from over 17 to 36 million tons
by Christopher Columbus during his travels after
as fresh. The area harvested followed a
the discovery of America in the fifteenth similar trend, with an increase of the surface
century and later spread to Africa and cultivated area of about 35% in the last 20
Asia. Early imported varieties belong to [Link], being today about 3.8 millions of
chinense (Scotch Bonnet or Habanero) which hectares. Fresh pep- per is cultivated in 126
most probably were the most consumed duringcountries of the world in all the continents. The
that time (Walsh and Hoot 2001). The world’s largest producer is China with over 18
flourishing commercial exchanges of Spanishmillion tons annually, fol- lowed by the Mexico
and Portuguese facilitated the spread of pepper
with about 3.5 million tons (FAOSTAT 2017).
around the globe, with an immediate successDry pepper is cultivated in 70 countries and no
due to a well acclimatization in the regions,
relevant production is reported in Oceania.
where they were used as a spice from that India is the largest producer with about 2.0
part of the population who could not afford to
million tons, followed by Thailand (349.615
purchase cinnamon, nutmeg, and other spicestons). Peppers are grown almost all over
that are widely used for seasoning and
the world and are fairly easy to cultivate both
preserving food. To date, the existence of in the field and in the greenhouse in a wide
35 range of climatic and environmental
Capsicum species is reported (Carrizo conditions. Africa, Europe, and America
García et al. 2016), five of which, namely, C.
contribute in the same proportion to the total
annuum, C. baccatum, C. chinense, C. world production (about 10–12% each) for
frutescens, and C. pubescens have beenfresh pepper; while for dry pepper, Asia and
domesticated and widespread with different Africa are the main pro- ducers contributing to
terms depending on the region of cultivation.
the 70.3 and 21.2%, respectively (Fig. 1.1).
In Mexico and Central America, the crop is The economic value of pepper production
called “chile” which was the ancient name has increased since 1991 becoming a good
given by local populations of the new world,
source of income for producers in many
in American English it becomes “chilli,” incountries and giving an important role in
Caribbean and countries Latin American international trading. The present worth of dry
countries it is commonly referred to as “ají” and
pepper is 3.8 billion dollars, while fresh
“rocoto,” from which derived names of many pepper contributes with 30,208 billion dollars.
cultivars of different species today present on
For both, the increase observed over the past
the market (i.e., aji Amarillo, aji limon, aji
25 years is four times higher in dry pepper
panca, rocoto manzano, rocoto brown, and and six times higher in fresh pepper.
rocoto de seda). It is also known as Around the genus Capsicum, there is an
pimiento (Spanish), red pepper and pepper increasing interest and fascination due to the
(English), pepper (Italian), piment (French),
amazing diversity in many characteristics, such
paprika (Ger- man and other northern European
as plant architecture, flower morphology,
languages). Overall, the present term “chili
fruit typology, colors, pungency, and qualitative
pepper” refers to varieties with small and spicy
traits which make this crop extremely
fruits, on the contrary, the term “sweet versatile and suitable for innumerable uses. As
pepper” refers to vari- eties with larger fruits
food, a variety of recipes are ensured thanks to
and little or no spicy. the presence of sweet and hot types. The
former are mainly widespread in temperate
regions of Europe and North America where
1.2 Economic and Culinary they are used freshly or cooked as vegeta- bles.
Importance The latter are instead mainly spread in the
tropical regions of America, Africa, and Asia,
World pepper production has grown where they are mostly consumed fresh or dried
consider- ably over 20 years (1997–2017, as condiment as spice in powder or salsa in
[Link]/ faostat), from 2 to about 4.5 many
million tons of dry
31 The Capsicum Crop: An P. Tripodi and S. 3

Fig. 1.1 Production share of dry and fresh pepper by region (FAOSTAT 2017)

Fig. 1.2 Examples of popular pod types of hot and sweet peppers. Photo credit Susan Lin, World Vegetable
Center, Taiwan

dishes. Food uses of peppers could then be whole fruits and powder (Poulos 1994). Based
sum- marized in the following classes: (a) fresh on pod shape and size, more than 20 market
use, of immature green fruits, mature red fruits, types (e.g. bell, cayenne, ancho, jalapeño,
and leaves; (b) fresh processing, for sauces, pasilla, Hun- garian wax, jwala, and Thai) are
pastes, pickles, beer etc.; (c) dried spices, commercially cultivated (Fig. 1.2).
from mature Furthermore, within each of
 41 The Capsicum Crop: An P. Tripodi and S. 4
these market types, there may be several literature. High contents of other essential
variants; for instance, bell may have blocky, vitamins such vitamin A in the form of b-
conical, or mini pods and cherry bell may have carotene and vitamins of group B
small or big pods (Fig. 1.2).

1.3 The Properties of

Pepper

The uniqueness of pepper is the typical

pungency due to the presence of
capsaicinoids. Capsaici- noids are secondary
metabolites and derivatives of
phenylpropanoids produced in placental
epider- mis cells and accumulated in structures
(blisters) located on the placenta surface
(Stewart et al.
2007). The hotness sensation when consumed
is given by the interaction with vanilloid
receptors, supposed to be a mechanism of
defense against mammalian herbivory.
Capsaicin and dihydro- capsaicin are the two
predominant compounds, accounting for almost
90% of total capsaicinoids. Anti-inflammatory,
anticancer, and anti-obesity activities have
been recognized within capsaici- noids (Luo et
al. 2010). These properties are exerted by
the release of substance P, a neuro-
transmitter involved in pain transmission by
nerve (Gamse et al. 1981). Peppers are also an
extremely good source of compounds
exerting antioxidant properties and responsible
for fruit pigmentation. Different colors are
encountered in mature fruits as a result of
accumulation of carotenoids in chro- moplasts
during ripening such as capsanthin and
capsorubin (mainly in red fruits), violaxanthin
and neoxanthin (mainly in yellow fruit), and
lutein and b-carotene (mainly in orange fruit)
(Gómez-Gar- cía and Ochoa-Alejo 2013).
Fruits are further well-known to have played a
leading role in the discovery of vitamin C by
Albert Szent-Györgyj, who extracted the first
pure chemical compound from Hungarian
paprika and was awarded Nobel Prize for
Medicine and Physiology in 1937 (http://
[Link]). Indeed, within
Capsicum species, a high level of ascorbic acid
(vitamin C) able to satisfy the recommended
daily intake (FDA
2018, attested to 60 mg for 100 g of raw
pepper) is commonly found in both sweet and
hot types and widely documented in the
 51 The Capsicum
(thiamine, riboflavin,
Crop: An and niacin) are P. Tripodi and S. 5
recognized. All these compounds, of which
content is deter- mined by species, cultivar,
environmental condi- tions, and maturation
stage, exert their biological effects protecting
cells against oxidative damage through the
interaction with oxygen molecules and
scavenging peroxyl radicals (Padayatty et
al.
2003; Howard and Wildman 2007). Finally,
antimicrobial and antivirulence properties,
against Streptococcus pyogenes, a major human
pathogen (Marini et al. 2015) and Fusarium
infection (Tewksbury et al. 2008) a
polyphagous fungus affecting many vegetables,
are reported. All these properties make pepper
a good candidate against diseases.
Other than food uses are recognized as
active ingredient in cosmetics,
pharmaceuticals, and pest management
(Bosland and Votava 1999). The extractable
colors from fruits due to the presence of
compounds unique in pepper such as capsanthin,
capsorubin, and cryptocapsin are extensively
used in the food processing industry as natural
colorant for a wide range of products such
meats, cheeses, and other foods. Non-food
uses include (a) coloring and flavoring
agents, from oleoresins (carotenoids) extracts
or powder, as example, paprika powder can be
used to inhibit lipid oxidation of pork meat
while oleo- resin is used to enhance physical
and sensory properties of food products
(Baenas et al. 2019); (b) ethno-
botanical/traditional medicine, from fruit
extracts and powders (pungent fruits); (c)
modern medicine/pharmaceuticals, from
extracts of capsaicinoids and carotenoids
which can exert analgesic, antimicrobial,
antioxidant, and anti-inflammatory effects; (d)
insecticides/re- pellents and antibacterial effect
from capsaici- noids extracts and organic acids
(i.e., cinnamic, coumaric, ferulic, and caffeic);
(e) spiritual, using whole fruits, e.g., “ristras”;
(f) ornamental, using whole plants or fruits;
(g) defense/punishment, using capsaicin
extracts/or powder (Kumar et al.
2006). The use in cosmesis is favored by
the presence of natural compounds which
allow to avoid allergies and other side
effects and are addressed to protect skin
oxidative and UVA-mediated damage
having thanks to the anti-wrinkle action and
fighting against free
 61 The Capsicum Crop: An P. Tripodi and S. 6
radicals (Baenas et al. 2019). The industrial C. praetermisssum; Pubescens which only
preparations are based on oleoresins rich of includes the homonymous domesticated
the above-mentioned bioactive compounds. species.
Finally, among the most curious aspects of the C. annuum is commercially most popular
Capsicum genus, there is certainly the rampant species worldwide. This species is
interest of many, searching and collecting, characterized by pungent and non-pungent
even in urban contexts, different species, accessions with herb or sub-shrub growth and
characterized by a wide variety traits, as well fruits having dif- ferent size, shape, and
as ornamental, aes- thetically appreciated or colors at maturity. C. frutescens and C.
rare varieties. This is evident in the rise of chinense are mainly culti- vated in American,
associations and websites dedicated to the Asian, and African countries. The former
subject. includes pungent accessions with fruits
predominantly small with less than 2 cm of
length, the latter instead comprise accessions
1.4 Genetic highly pungent and irregular shape of fruits.
Resources and The other two domesticated species (C.
Breeding baccatum and C. pubescens) are cultivated in
Central and South America and are
The Capsicum genome has an estimated size distinguished by particular phenotypic
of characteristics such as the yellow or green
3.5 Gb and includes mainly diploid species spots in the corolla (C. baccatum) or the dark
with colored seeds (C. pubescens). Several wild
12 chromosome (2n = 2x = 24). Within species are part of the genus Capsicum and
the genus, there are also recognized species are principally distributed in the area of
with 13 chromosomes (2n = 2x = 26) as origin (Table 1.1). All of them are
well as one tetraploid species (2n = 4x = characterized by very small oval or spherical
48) which is C. annuum var. glabriusculum, fruits (Fig. 1.3) with specific distinctive traits
the wild form of the cultivated pepper. Recent related to flower color (white, yellow, and
investigations have grouped the Capsicum purple with different type of spots), seed color
species in 11 clades according to main (brownish or black), and flower shape (stellate,
morphological features, provenance, and rotate, or campanulate) (Barboza and Bianchetti
phylogenetic relationships (Carrizo García et al. 2005). Although the uniqueness and beautiness
2016) (Table 1.1). The spe- cies of greatest distinguish many species of pep- per, most of
interest for consumption and breeding are in the breeding activities have been carried out
three main clades namely: Annuum which within the Annuum clade due to the lack of
includes three domesticated (C. annuum, C. interspecific barriers between C. annuum, C.
frutescens, and C. chinense) and two wilds chinense, and C. frutescens (Pickersgill 1997;
(C. annuum var. glabriusculum and C. galapa- Perry et al. 2007). However, the
goense); Baccatum including three forms incompatibility occurring across clades could
of C. baccatum (var. baccatum, var. pendulum, be overcome using aids such as embryo
and var. umbilicatum) and the wilds C. rescue. Wild and
chacoense and

Fig. 1.3 Mature fruits of wild Capsicum species: a C. chacoense, b C. praetermissum, c C. eximium, d C. annuum var.
 71 The Capsicum
glabriusculum, and eCrop:
C. flexuosum
An P. Tripodi and S. 7
81 The Capsicum Crop: An P. Tripodi and S. 8
Table 1.1 Capsicum clades and related species, main features and native area
Cladea, species name, Pungency Fruit colorb Area of origina
chromosome number
1. Annuum (x=12)
C. annuum Non-pungent Variable Central and south America regions
and pungent
C. annuum var. Pungent Red Venezuela, central america
glabriusculum
C. chinense Pungent Variable Central America, Colombia, Ecuador,
south-eastern Brazil, Venezuela
C. frutescens Pungent Variable Central America, central-eastern Brazil,
Colombia, Ecuador,Venezuela
C. galapagoense Pungent Red Galapagos Islands
2. Baccatum (x=12)
C. baccatum var. baccatum Non-pungent Variable Argentina, Bolivia Paraguay, Peru’
and pungent
C. baccatum var. pendulum Non-pungent Variable Argentina, Bolivia Paraguay, Peru’
and pungent
C. baccatum var. pungent Variable Argentina (north and central), Bolivia
umbilicatum (lowlands)
C. chacoense Pungent Red Argentina, Bolivia, paraguay
C. praetermissum Pungent Red South-eastern Brazil
3. Tovarii (x=12) Pungent
C. tovarii Pungent Red Perù
4. Pubescens (x=12) Pungent
C. pubescens Pungent Variable Argentina, Bolivia, central America,
Ecuador, Perù
5. Purple corolla (x=12) Pungent
C. cardenasii Pungent Red Bolivia (highlands)
C. eximium Pungent Red Argentina (north and central), Bolivia
(lowlands)
C. eshbaughii* Pungent Red Bolivia (lowlands)
6. Atlantic forest (x=13) Pungent
C. campylopodium Pungent Greenish-yellow South-eastern Brazil
C. cornutum Pungent Greenish-yellow South-eastern Brazil
C. friburgense Pungent Greenish-yellow South-eastern Brazil
C. hunzikerianum Pungent Greenish-yellow South-eastern Brazil
C. mirabile Pungent Greenish-yellow South-eastern Brazil
C. pereirae Pungent Greenish-yellow South-eastern Brazil
C. recurvatum Pungent Greenish-yellow South-eastern Brazil
C. schottianum Pungent Greenish-yellow South-eastern Brazil
C. villosum var. villosum Pungent Greenish-yellow South-eastern Brazil
(continued)
91 The Capsicum Crop: An P. Tripodi and S. 9
Table 1.1 (continued)
Cladea, species name, Pungency Fruit colorb Area of origina
chromosome number
7. Longidentatum (x=13)
C. longidentatum Non-pungent Greenish-yellow Central-eastern Brazil
8. Bolivian (x=*)
C. caballeroi Pungent Red Bolivia (lowlands)
C. minutiflorum Pungent Red Bolivia (lowlands)
C. ceratocalyx Pungent Red Bolivia (highlands)
C. coccineum Pungent Red Bolivia, western Brazil
9. Flexuosum
C. flexuosum Non-pungent Red South-eastern Brazil, north-eastern Argentina
and pungent and eastern Paraguay
10. Caatinga (x=13)
C. caatingae Pungent Greenish-yellow Central-eastern Brazil
C. parvifolium Pungent Greenish-yellow Central-eastern Brazil, Colombia, Venezuela
11. Andean (x = 12)
C. rhomboideum Non-pungent Red Central America, Colombia, Ecuador, Perù,
Venezuela
C. scolnikianum Non-pungent Red Colombia, Ecuador, Perù
C. geminifolium Non-pungent Red Ecuador, Perù
C. lanceolatum Non-pungent Red Ecuador, Perù, central America
C. dimorphum Non-pungent Red Colombia, Ecuador, Perù
a
according to Carrizo García et al. (2016)
b
at maturity stage
*chromosome number not reported

domesticated species have been used Capsicum genetic resources including the
particularly for disease resistance and results pro- gress in breeding and genomics are
are widely documented in the literature. discussed further in the chapters to be
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[Link]/nobel_prizes/medicine/laureates/