Scientia Horticulturae 288 (2021) 110306

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Scientia Horticulturae
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Review

Tomatillo or husk tomato (Physalis philadelphica and Physalis ixocarpa):

A review
Julio Emmanuel González-Pérez, José Ángel Guerrero-Beltrán *
Departamento de Ingeniería Química, Alimentos y Ambiental, Universidad de las Américas Puebla. Ex hacienda de Santa Catarina Mártir, San Andrés Cholula, Puebla
72810, Mexico

A R T I C L E I N F O A B S T R A C T

Keywords: Tomatillo (Physalis philadelphica and Physalis ixocarpa) are native fruits to Mexico and Central America. However,
Fruit origin the tomatillo plant grows in tropical and subtropical regions around the world. The tomatillo fruit is also called
Plant husk tomato. The growth conditions may modify, as for another vegetable, the yield and composition of fruits
Botanical aspects
and seeds. Pests and some microorganisms (transmitted by insects) generate some diseases to the plant. Plants
Chemical composition
and fruits of tomatillo contains vitamins, minerals, phenolic compounds, and steroidal lactones such as physalins,
Nutritional compounds
Health benefits having antimicrobials and antinarcotic effects. This is why plants and fruits are widely used in traditional
medicine to relieve some discomforts: fever, cough, amygdalitis, gastrointestinal disorders or diabetes and as
food ingredient in the Mexican and Guatemalan cuisines. The aim of this review was to compile in a single
manuscript the whole information about the plant and fruits of tomatillo, considering fruit origin, botanical
aspects, cultivation characteristics, components, and health benefits as well as the main uses of fruits and parts of
the plant.

1. Introduction 2019). Other studies have developed different technical procedures to

Tomatillo belongs to the genus Physalis (Greek for “a bladder”)
(Robledo-Torres et al., 2011). Tomatillo is an annual plant original to
Mexico and Guatemala. The plants growth widely in tropical and sub­
tropical regions all over the world, predominantly in some countries of
the American continent (Small, 2012; Zhang et al., 2016). Tomatillo
fruits are of green, yellow, purple, and purple-green colors, covered with
a thin husk (calyx) (Small, 2012; Whitson, 2016). Fruits and plants of
Physalis philadelphica and Physalis ixocarpa species are nutritionally and
commercially important (Pretz and Deanna, 2020; Zhang et al., 2016) in
several countries. Plants and fruits of tomatillo contains vitamins,
minerals, phenolic compounds, secondary metabolites such as physalins
and steroids (steroidal lactones). The plant and fruits are widely used in
traditional medicine to relieve some ailments (relieve fever, cough,
amygdalitis, gastrointestinal disorders or diabetes) and as food ingre­
dient in the Mexican and Guatemalan cuisines (García-Mendieta et al.,
2012; González-Chavira et al., 2019; Maldonado et al., 2011).
Some studies have analyzed the characteristics of the main problems
found throughout the growth of tomatillo plants. Different microor­
ganisms and insects are responsible of various diseases (Sastry et al.,
increase the yield of fruits and components in the fruit throughout the
entire germination process, growing of the plant and development of its
parts (García-Osuna et al., 2015; Marín-Sánchez et al., 2007; Martí­
nez-Solís et al., 2006; Montejo-Canul et al., 2019; Peña-Lomelí et al.,
2018; Robledo-Torres et al., 2011).
Accordingly, the aim of this review was to compile information about
the plant and fruits of tomatillo; considering the fruit origin, botanical
aspects, cultivation characteristics and their components; in addition to
the main uses of the fruit and different parts of the plant.
2. Fruit origin and botanical aspects
2.1. Names
Tomatillo fruit (husk tomato) is produced by the plants of
P. philadelphica Lam. and its subspecies P. ixocarpa Brot. (Sobrino-Ves­
perinas and Sanz-Elorza, 2007). Table 1 shows their main differences. To
clarify the taxonomic classification of this species, some studies
analyzed cytological and taxonomic characteristics. One of the first
differences found was that the peduncle of P. ixocarpa is shorter than

* Corresponding author.
E-mail address: angel.guerrero@udlap.mx (J.Á. Guerrero-Beltrán).

https://doi.org/10.1016/j.scienta.2021.110306
Received 7 January 2021; Received in revised form 13 April 2021; Accepted 19 May 2021
Available online 11 June 2021
0304-4238/© 2021 Elsevier B.V. All rights reserved.
J.E. González-Pérez and J.Á. Guerrero-Beltrán Scientia Horticulturae 288 (2021) 110306

Table 1 “tomate de capote” (in Colima, Mexico), tomate (in central Mexico),
Main difference between Physalis philadelphica and Physalis ixocarpa. “tomate milpa”, “miltomate” (in Oaxaca, Mexico and Guatemala),
Characteristics Physalis philadelphica Physalis ixocarpa Mayan husk tomato, ground cherry, and “jamberry” (introduced by
a Iowa State College) (Gollapudi and Motohashi, 2013; Jiménez-Santana
Plant size 15–60 cm 60–120 cmb
Leaves size – 6.25 cm in long and 3 et al., 2012).
cm widec
Corolla size 0.8–3.2 cm in diametera 1.25 cm in long and 2 2.2. Origin
cm wideb
Size fruit 1.5 cm in diameter (wild)d
3–7 cm in diameter (cultivated) Small (<4.5 cm in
archeological excavations in Tehuacan Valley (Mexico) suggest that
diameter)e native people, in their diet, incorporated “tomatillos” since 900 years
Pedicel lengthg Large (0.05–0.10 cm) a Short BC. Aztec and Mayan cultures domesticated P. ixocarpa Brot. (Fig. 1).
Locatione,g Mexico, Guatemala, USA and Mexico (Southern) and There is evidence of wild growth of tomatillo from California in the
Southern parts of Canada Guatemala
United States to Guatemala and Nicaragua (Jiménez-Santana et al.,
Domesticationg Wild form Domesticated
Taxonomic 1786 1819 2012). Wild tomatillo seeds were found in archeological sites from Great
description yearh Plains-Eastern United States such as winter camps of the Fort Ancient
Composition (fresh)i (1000–1750 AD, in east-central Kentucky), Dymock sites (1000–1200
β-carotene 21 µg/100 mg 2 µg/100 mg AD, in Ontario), at Meehan-Schell site (850–1100 AD, in Iowa)
Lutein 12 µg/100 mg 1 µg/100 mg
(Kindscher et al., 2012). Tomatillo is cultivated in tropical and sub­
a
Hernández and León (1994). tropical areas of the world (Small, 2012).
b
Shenstone et al. (2020). 2.3. Characteristics of flowers
c
Duarte and Paull (2015).
P. philadelphica Lam. and P. ixocarpa Brot. are members of the Sol­
d
Zamora-Tavares et al. (2015).
anaceae family (Robledo-Torres et al., 2011). Both species are
confounded between them or are considered synonymous species
e
Robledo-Torres et al. (2011); because some varieties are diploids (2n = 2x = 24), have blue anthers
f
Menzel (1951). that twist after dehiscence (Fig. 2A-2B), a yellow corolla with five
smudges (Fig. 2B), and a ten-ribbed calyx attached as a “bag” (Fig. 2D).
g
Pretz and Deanna (2020). The calyx or husk shields the tomatillo fruit (Gollapudi and Motohashi,
2013; Kindscher et al., 2012; Pretz and Deanna, 2020; Whitson, 2016).
h
Sobrino-Vesperinas and Sanz-Elorza (2007).

i
2.4. Fruit characteristics
Elizalde-González and Hernández-Ogarcí (2007).

Tomatillo fruits are small, spherical or slightly oblate (Tables 1 and

that of P. philadelphica (Menzel, 1951; Pretz and Deanna, 2020).
Waterfall (1967) found that in some Mexican and Central American
species, the corolla size of P. ixocarpa (approximately 2 cm in diameter)
is smaller than that of P. philadelphica (3.2 cm in diameter). However,
other authors have not reported significant differences between the
corolla sizes of these species (Hernández and León, 1994; Shenstone
et al., 2020). Fernandes (1974), based in cytological evidences,
distinctive stigma and small flowers, concluded that P. ixocarpa and P
philadelphica are distinct species. On the other hand, some researchers
(Vergara-Ponce et al., 2011; Zamora-Tavares et al., 2015; Sando­
val-Padilla et al., 2019) have worked in elucidating differences of
several tomatillo species using different genetic techniques. For
instance, Vergara-Ponce et al. (2011) used six ISSR (Inter-Simple
Sequence Repeats) primers to determine their usefulness for taxonomic
discrimination and elucidate their use for inferring the interspecific
relationship among Physalis virginiana Miller, Physalis cordata Miller,
Physalis cinerascens (Dunal) Hitch, Physalis lignescens Waterfall, Physalis
sulphurea Waterfall, Physalis angulata L., Physalis lagascae Roem. &
Schult. and Physalis philadelphica Lam. They found that the taxonomic
discrimination by ISSR produced different fingerprint profiles and that
the ISSR markers estimate genetic relationship in Physalis species.
Tomatillo fruits are popularly known as “tomate” (no the red one),
“tomatillo” (“small tomato”) in Mexico and Central America (Pretz and
Deanna, 2020; Sobrino-Vesperinas and Sanz-Elorza, 2007) or husk to­
mato (in English). The tomate word is traced to the Nahuatl word
“tomatl”, which refer to a globose fruit with many seeds and watery flesh
(Small, 2012). However, some reports indicate that the word correspond
to native terms in Mesoamerican language, specifically proto-Zapotecan
(3350 BC) (Brown, 2010). Other names are “tomate verde” (in the
Northern states of Mexico), “tomate cascara”, “tomate milpero” (in the
Southern states of Mexico), “fresadillas” (in Nuevo Leon, Mexico),
2) and surrounded by an enlarged calyx (Smith et al., 1999)
(Figs. 2D-2F). In general, the fruit size has been reported between
27.42–54.57 mm in equatorial diameter and 25–44.32 mm in polar
diameter (Table 2). The species "CHF1 Chapingo", "Commercial", "L171",
"L17", "L182", "L94", "L97", "L30" and "Feira de Santana" have an equa­
torial diameter 16% larger than the polar diameter. On the hand, some
species can have similar size in diameter as “Felipe Ángeles”, “Gran
Esmeralda”, “Purple Tamazula” or “Rendidora”. The pericarp color of
mature fruits is usually green, yellow, bright orange-red, purple, or
purple-green (Gollapudi and Motohashi, 2013). The pulp is pale-yellow,
crisp or soft, acid, sweet with characteristic flavor, and has many tiny
seeds (Fig. 2F) (Duarte and Paull, 2015; Small, 2012). The varieties with
green pericarp are “Rendidora”, “Gigante”, “Tamayo”, “Toma Verde”,
“Gulliver hybrid”, “Manzano”, “Milpero”, “Salamanca”, “Tamazula”,
“Silvestre”, “Arandas”, and “Puebla” (Escobar-Guzmán et al., 2009;
Mulato-Brito and Peña-Lomelí, 2007). The varieties with purple pericarp
are “Purple coban”, “Purple de milpa”, “Purple hybrid” and “Plum jam”
(Slivoviy Jam) (Naumova et al., 2019).
3. Cultivation and harvest
3.1. Production
Fig. 3 shows the average production of tomatillo in Mexico between
2015 and 2019. Total production was 754,013 tons/year (SIAP, 2020).
Mexican areas with the highest average seasonal production in the
spring-summer cycle (2015–2019) were Zacatecas (90,659 tons), Jalisco
(52,444 tons), Puebla (36,463 tons), Mexico State (30,322 tons),
Michoacán (24,571 tons), Morelos (20,820 tons), Tlaxcala (20,806
tons), and Hidalgo (15,121 tons). In the fall-winter cycle (2015–2019),
the main producers were Sinaloa (142,822 tons), Sonora (35,523 tons),
Nayarit (33,426 tons), Michoacán (25,996 tons), Jalisco (20,091 tons),

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J.E. González-Pérez and J.Á. Guerrero-Beltrán
Fig. 1. Origin of tomatillo.
Puebla (18,379 tons), Chiapas (14,972 tons), and Guanajuato (13,186
tons) states in Mexico.
3.2. Plant characteristics
Tomatillo crop is grown best at low altitudes, between 18 and 25 ◦ C,
slightly acid soils, and areas with 15–120 cm annual rainfall (Hernández
and León, 1994; Shenstone et al., 2020). The tomatillo plant life cycle is
short (3–4 months) (Moriconi et al., 1990). The growing time takes
about 12–14 weeks. During this time, occurs characters such as begin­
ning of flowering, opening, and beginning of harvest (Mulato-Brito and
Peña-Lomelí, 2007).
The tomatillo plants have smooth branches and leaves (Fig. 2A).
Their leaves (6.25 cm long and 3 cm wide) are ovate, pointed (at the
apex), wedge-shaped (at base), with wavy margins (in some varieties).
Flowers of tomatillo plant sprout singly in the leaf axils and have a fused
five-toothed green calyx (Fig. 2B). The petals are yellow with dark
brown spots in the throat. The flower size is about 0.8–3.2 cm in length
and 2 cm in wide. During flowering, the calyx enlarges for cover the fruit
(Duarte and Paull, 2015). The calyx helps to protect the fruit from in­
sects, birds, diseases and extreme climate conditions like heavy rain,
hailstorms, extreme temperature changes or radiation (Cruz-Álvarez
et al., 2012).
Tomatillo fruit can be harvested at various development stages
(mature or immature). For commercial marketing, the fruit is harvested
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Scientia Horticulturae 288 (2021) 110306
when it reached its maximum size. In other words, when the calyx is
completely filled or broken by the fruit (Cantwell et al., 1992; Smith
et al., 1999).
Cruz-Álvarez et al. (2012) analyzed the influence of calyces on to­
matillo after postharvest and stored at 4 and 20 ◦ C. The fruits with
calyces stored for 20 days after harvest had the highest content of
chlorophyll (0.147–0.161 µg/100 g) than tomatillos without calyces.
This was related to the degradation of chlorophyll by the light. On the
other hand, the presence of calyces generated greater mass reduction
due to the water loss by transpiration and low relative humidity.
Tomatillos (with or without calyces) stored at 4 ◦ C maintained (p <
0.05) their firmness and total soluble solids or ◦ Brix (%) content, while
fruits stored at 20 ◦ C showed 5% weight losses.
3.3. Cultivation practices
Direct seeding and transplantation are common cultivation practices
of the Solanaceae plants. The first one consists in the spreading of seeds
directly into gardens or fields. This procedure is limited by low tem­
peratures because the plant growth is poor at temperatures below 18 ◦ C.
Seeds are viable to germinate within three years (Hernández and León,
1994). Commonly, seeds germinate in about 7–10 days; afterward, a
primary shoot elongation occurs along four weeks. Before the primary
shoot elongation occurs, the first flower buds are formed (in about 3–4
weeks), which continues until the plant senescence (in about 14–15
J.E. González-Pérez and J.Á. Guerrero-Beltrán Scientia Horticulturae 288 (2021) 110306

Fig. 2. Tomatillo (Physalis ixocarpa) diploid variety. A) P. ixocarpa Brot., B) Flower bud open, C) Flower bud closed, D) Tomatillo fruit with calyx, E) Tomatillo
without calyx, F) Total fruit content.

Table 2
Fruits characteristics of some Physalis ixocarpa varieties.
Variety Origin Weight (g) Diameter (mm) ◦
Brix (%) pH References
Equatorial Polar

Felipe Ángeles (Landraces) Acatzingo, Puebla 34.08–59.90 38.43–54.57 35.50–44.32 5.61–6.45 3.85–4.17 Ramírez-Godina et al. (2013)
Gran Esmeralda Harris Moran 32.51–54.99 36.26–51.32 34.63–42.21 5.48–6.42 3.80–4.06
Purple Tamazula (Landraces) Arandas, Jalisco 13.28–18.84 27.42–36.18 26.43–30.87 5.55–6.89 3.99–4.29
Rendidora (diploid) Zacatepec, Morelos 26.75–35.59 34.56–43.30 32.43–35.85 5.81–6.45 3.79–4.15
Tecozautla Santa Cruz Xoxocotlán, Oaxaca 52.29 49.0 Benito-Bautista et al. (2015)
Rendidora 26.10 38.4
Diamante 35.19 42.4
San Martín 37.29 42.2
CHF1 Chapingo Texcoco, Mexico State 26.4 45.4 34.1 Ponce-Valerio et al. (2011)
Commercial (purple) Brazil 13.13 30.33 26.12 2.33 4.05 Curi et al. (2017)
L171 (purple) Morelos, Mexico 53.5 49.0 42 González-Chaviraet al. (2019)
L17 (purple) Guerrero, Mexico 14.3 31 25
L182 (green) Morelos, Mexico 60.6 54 40
L94 (green) 34.9 44 33
L97 (green) 28.4 38 31
L30 (green) Puebla, Mexico 16.0 33 26
Feira de Santana (green) Brazil 38.3 44 35.9 5.0 Barroso et al. (2017)

weeks). After flowering (4–5 weeks) and a week later, the first fruits cultivation practices depends on the environmental characteristics, the
appear, reaching their maximum size after 8 weeks (Moriconi et al., presence of pest or consumers demand.
1990). Transplanting consist in the previous seed germination and
growing the crop indoors before transplanting outdoors (Hernández and
León, 1994). Some advantages of this procedure are the saving of seeds, 3.4. Pests and diseases of tomatillo plants
reduction of weeds and the possibility of starting the growing cycle
when the outside temperature is below 18 ◦ C. Finally, the choice of 3.4.1. Pests
The tomatillo plants can be infested by the beetle Epitrix cucumeris,

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J.E. González-Pérez and J.Á. Guerrero-Beltrán
Fig. 3. Average cycle production in Mexico of tomatillo between 2015 and 2019. Adapted from SIAP (2020).
Harris, the American serpentine leaf miner (Liriomyza trifolii, Burgess), a
fly, and the fruit worm (Chloridea subflexa, Guenée), a moth, formerly
called Heliothis subflexa (Montejo-Canul et al., 2019). H. subflexa is an
insect, which larva consumes the fruits of Physalis species (Bautista-­
Martínez et al., 2015). The insect uses the inflated calyx to protect itself
from predators. The presence of the insect in tomatillo reduce the fruit
quality (Oppenheim and Gould, 2002). Tomatillo is also host of Bac­
tericera cockerelli (Sulc), a fly. B. cockerelli injects a toxin to the plant,
which causes chlorosis; in addition, the insect can transmit a phyto­
plasma that generate a yellowish disease (Crespo-Herrera et al., 2012).
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Scientia Horticulturae 288 (2021) 110306
Therefore, growers of tomatillo should be alert about any infestation by
insects that may reduce the production.
3.4.2. Diseases of tomatillo plants
The main symptoms of tomatillo diseases include wilting, poor
growth, discoloration (vascular tissues), root rotting and the death of
plants (Ayala-Armenta et al., 2020). Table 3 shows some diseases of
tomatillo plants. Plants diseases are related to pathogens such as bac­
teria (Rivera-Martínez et al., 2017), molds (Ayala-Armenta et al., 2020;
Félix-Gastélum et al., 2007), and viruses (Adkins et al., 2006; Can, 1994;
J.E. González-Pérez and J.Á. Guerrero-Beltrán Scientia Horticulturae 288 (2021) 110306

Table 3
Microorganism and diseases of Physalis ixocarpa and P. philadelphica plants.
Species Country Disease Microorganisms Symptoms References

P. ixocarpa Mexico Tomatillo wilt Fusarium oxysporum, Neocosmospora solani Exterior part: yellowing of the foliage, Ayala-Armenta et al.
Brot. (syn. Fusarium solani), Rhizoctonia solani, growth reduced, wilting and plant (2020)
Pythium spp., Macrophomina phaseolina and death. Roots: dark brown rot (from neck
Sclerotium rolfsii until base of the stem).
Fruit: reduction of number of fruits/
plant and size
Impatiens necrotic Orthotospovirus (virus transmitted by Dark necrotic spots on leaves and stems González-Pacheco and
spot virus (INSV) transportation vectors) Silva-Rosales (2013)
Yellow mottle Alfalfa mosaic virus or Alfamovirus (AMV) Yellow mottle, mosaic, leaf deformation, De la Torre-Almaráz
(transmitted by insect Myzus persicae) chlorosis and wilt symptoms et al. (2003)
Leaf spots Cercospora spp. Dark brown lesions on leaves Félix-Gastélum et al.
(2007)
Powdery mildew Podosphaera xanthii Concentric circles on leaves or calyx
Mexico and Tomato yellow leaf Virus (transmitted by insect Bemisia tabaci) Chlorotic margins, yellowing and Gámez-Jiménez et al.
Guatemala curl virus (TYLCV) interveinal yellowing (2009); Salati et al.
(2010)
Mexico, USA Floral abortion and Candidatus liberibacter solanacearum, The plant ages prematurely and its yields Rivera-Martínez et al.
and Central yellowing (transmitted by insect B. cockerelli Sulc) decrease (2017)
America
Portugal Tomato chlorosis Virus (transmitted by insects Bemisia tabaci, Plants exhibited intermittent yellowing Trenado et al. (2007)
virus (ToCV) Trialeurodes vaporariorum and T. abutilonea)
USA (Louisiana) Physalis mottle virus Virus (transmitted by beetle Epitrix The leaves of plant exhibit mosaic and Can (1994)
(PhyMV) cucumeris Harris) yellow mottle symptoms
P. philadelphica Mexico golden Pepper (transmitted Virus mosaic, leaf curl whitefly) Yellowing Lozano and branchy top Méndez-et al. (2001)
by mosaic virus
(PepGMV)
Pepper Huasteco Virus (transmitted by insect Bemisia tabaci) Yellowing mosaic, leaf curl and branchy Valverde et al. (1993)
yellow vein virus top
(PHYVV)
USA (Georgia Tomato spotted wilt Virus (transmitted by insects Frankliniella Chlorosis and stunting of young leaves Adkins et al. (2006);
And Florida) virus (TSMV) occidentalis and F. bispinosa) and distinct chlorotic rings and rings Díaz-Pérez and Pappu
patterns on older leaves (2000)
USA Turnip mosaic virus Virus (transmitted by insect Myzus persicae) Several stunting and mosaic on leaves. Liu et al. (2012)
(California) (TuMV) Mottled and unmarketable on the fruit

De la Torre-Almaráz et al., 2003; Gámez-Jiménez et al., 2009;
González-Pacheco and Silva-Rosales, 2013; Liu et al., 2012;
Méndez-Lozano et al., 2001; Trenado et al., 2007; Valverde et al., 1993),
or phytoplasma (Santos-Cervantes et al., 2007). Hence, soils or any other
mean for planting tomatillo should have nutrients for avoiding or
reducing diseases.
3.5. Methods to increase yield of tomatillo
3.5.1. Plant propagation
3.5.1.1. Fertilization. Some of the most common fertilizers are nitrogen
(N), phosphorus (P), and potassium (K) (Smith et al., 1999). Nitrogen
participates in amino acids and proteins formation in the plant, pro­
motes leaves and stems growth, provides dark color to leaves, and in­
crease flowering, fruit setting and fruit development. Phosphorus is
essential for cell division, meristematic tissue development, fruit set and
growth, and ripening. Potassium regulates water consumption and
transpiration (due to water retention in cellular tissue), increases the
plants resistance to extreme temperatures, and increases the fruits size
(Briccoli et al., 2012; Devi et al., 2020).
One fertilization technique is to spread the fertilizer on the soil. Some
researchers have reported the minimum composition of fertilizers
spread on the soil for the tomatillo plant growth. 112–168 kg N/ha are
used in a typical fertilization, 44.8 and 89.6 kg phosphorus pentoxide
(P2O5)/ha have been used in early spring planting or in alkaline soils for
phosphorus enrichment, respectively, and 56–168 kg K2O/ha are used to
provide K to soils (Smith et al., 1999). Some fertilizers can be supplied
by roots. This technique consists of providing small quantities of min­
erals in assimilable form through the roots (Devi et al., 2020). The
objective of the technique is to increase yield and to improve quality of
the fruits (Trejo-Téllez et al., 2007). Trejo-Téllez et al. (2007) evaluated
the use of a combination of soil fertilization (N, P and K) with foliar
application for P. ixocarpa Brot. var. Rendidora growth. The composition
of the foliar fertilizer was 0.2 g CO(NH2)2/L, 0.1 g NH4NO3/L, 0.2 g
KNO3/L, 0.2 g KH2PO4/L, 0.05 g H3BO3/L, 0.05 g MnSO•44H2O/L, 0.05 g
CuSO•45H2O/L, 0.05 g ZnSO•47H2O /L, and 0.10 g Fe-EDTA/L. The
fertilization was carried out considering one part of soil fertilization and
one part of foliar application, once a week for 12 weeks. The soil used
was a fertile soil with neutral pH and the procedure was carried out
under protected cultivation. It was found that this method compensate
the nutritional deficiencies of N, Fe, Zn, Mn and B as well.
The use of an appropriate fertilizer at right time (early spring
plantings) or according the soils characteristics (alkaline soils) may help
to increase efficiency and reduce the cost of fertilization. The foliar
fertilization is a good alternative to reduce costs because it reduces the
amount of fertilizer used, the nutrients are given at the moment of
greatest need and is effective, in a short times, because increases the
absorption of nutrients.
3.5.1.2. Addition of solutions of salts. Environmental stress conditions of
plants have been tried to increase the level of phenolic compounds in
tomatillo fruits. Rosas-Medina et al. (2020) cultivated P. ixocarpa plants;
the crop was grown in a protected cultivation until fructification. The
plants were subjected to saline treatment for 15 days; it started 42 days
after sowing (15 days before the fruit develops). In addition, the treat­
ment was completed with 90 or 120 mM NaCl; results revealed that
plants treated with 120 mM NaCl increased the flavonoids and antho­
cyanins contents in the fruits by 1.16 and 1.46 times than in control,
respectively.
3.5.1.3. Plant treatment with bio-agents. Likhtaryk and Ananasovyi va­
rieties of tomatillo plant were treated with bio-agents (Humisol, Rost­
moment, Azotobacterin, Biomag, Biopolycid, and Phosphoretherin). The

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J.E. González-Pérez and J.Á. Guerrero-Beltrán
treatments consisted in spraying 5 times (10–12 days each) the plant
with the bio-agent. The plants without treatment had heights in the
range 59.2–61.1 cm in Likhtaryk variety and 56.9–62.7 cm in Anana­
sovyi variety. The fruits of both varieties without bio-agent presented
2.14 cm in diameter and 4.9–5.4 g in weight. The fruits treated with bio-
agent had 2.5–2.6 cm in diameter and 5.2–6.2 g in weight. The use of
Humisol and Biomag sprays increased the height of the plant in 2 and
10%, respectively. The Biomag agent accelerated the budding process
and the fruits formation (Vdovenko et al., 2018).
3.5.2. Seed germination
The seed germination is an important characteristic for the tomatillo
crop production (Sánchez-Vásquez et al., 2007). Martínez-Solís et al.
(2006) evaluated the temperature (20, 25, 30 and 35 ◦ C) effect on the
germination of P. ixocarpa seeds. The germination test was done with
seeds of 16 different tomatillo varieties (Manzano, Tamazula-Erecta,
Orizaba, Pob-8–201-SCM-H-96-S527, Fam130, Selección-Morelos,
Tamazula-Rastrera, Salamanca, Selección-Tecamac, Sintético-
Intervarietal-Pue × Chap., Selección-Masal-SJ-27, CHF1-Selección-
Masal, Población-FMHM, Miltomate-Morado-Amarillo, Sintético-96-
Interfam.-Santa-Lucía, Verde-Puebla). The authors found that the
optimal temperature of germination of tomatillo seeds was 30 and 35 ◦ C.
That temperature range helps to reach a seeds germination percentage of
65.85–67.29% and an increase in the germination rate of
23.98–25.69%.
Another method to accelerate the germination of tomatillo seeds is
the “osmopriming”. It consists of immersing the seeds in osmotic solu­
tions of mannitol, polyethylene glycol and NaCl through a given period.
In this condition, the seeds maintain a germinate state during a
controlled rehydration (between the hydric potential of the solution and
interior of the seed). As a result, seeds may reduce their germination
time. (Marín-Sánchez et al., 2007; Mouradi et al., 2016). Marín-Sánchez
et al. (2007) evaluated the effect of osmopriming on the germination of
tomatillo seeds of “Rendidora” variety with KNO3, polyethylene glycol
and KCl as osmotic agent at − 5 to − 20 atm during 48 to 96 h. Seeds
reached a germination time of 252 h and a germination percentage of
66.50%. The use of the osmotic system at − 5 to − 20 atm in osmoprimed
seeds reduced 6–13% the germination time and increased the germi­
nation percentage in 25.1–31.6%; the immersion time did not modify
the percentage of germination (p > 0.05). The best osmopriming con­
ditions of Rendidora seeds were using KNO3 as the osmotic agent at − 15
atm/48 h and − 20 atm/48 h. Those conditions gave germination per­
centages of 87.5 and 90.5% respectively.
3.5.3. Pruning
Pruning is a technique used to regulate the balance between vege­
tative and reproductive growth. This technique helps to control the
growth of plants and the quality of the fruits. It consists in removing
parts of a plant for inducing the motion of reserve substances and in­
crease water nutrients (Hernández et al., 1992). Ponce-Valerio et al.
(2011) analyzed the effect of four pruning levels (fourth-inter-node,
sixth-inter-node, eight-inter-node and without pruning) on two
P. ixocarpa Brot. ex Horm. varieties. The varieties analyzed were CHF1
Chapingo and Tamazula SM2. Authors found that only the
fourth-inter-node cutting was essential to obtain a high total yield. The
CHF1 Chapingo variety had the highest yield (963.49 g/plant) and fruit
quality (26.4 g in weight, 45.4 mm in equatorial diameter).
3.5.4. Protected cultivation
Protected cultivation was implemented to increase yields of toma­
tillo fruits of Rendidora variety. This technique was carried out under
three protective structures covered with sheets of different materials.
The sheets of the structures were constructed using transparent poly­
ethylene, diffuse white polyethylene and white mesh. Plants grown in
the open field received 51.02 mole/m2/d of photosynthetically active
radiation (PAR). Transparent polyethylene, diffuse white polyethylene
7
Scientia Horticulturae 288 (2021) 110306
and white mesh sheets transmitted 35.58, 25.62 and 38.19 mole/m2/d,
respectively, to plants. The use of transparent polyethylene sheets
increased the temperature of the system (18.01–22.17 ◦ C) compared to
open field system (16.86–20.25 ◦ C). In addition, that sheet materials
increased the water consumption (84 L/plant) and yield (3.23 kg/plant)
(Ramos-López et al., 2017).
3.5.5. Pollination by hand
Wind or insects perform natural pollination of tomatillo. Fig. 2.C
shows a flower closed, it occurs when the flower has been pollinated,
after that, the flowers do not reopen, they get whiter and then fall down.
The tomatillo seeds were considered mature after 55 days of flowering
(Pérez-Camacho et al., 2012). However, this process does not occur
naturally in a greenhouse; therefore, pollination by hand is commonly
used to achieve the greatest fruit size and seeds production. Peña-Lomelí
et al. (2018) determined the optimal time to carry out pollination by
hand of Diamante (hybrid of CHF1 Chapingo and Puebla SM3 varieties)
and Manzano Tepetlixpa varieties. The authors found that the optimal
period is between two (Manzano Tepetlixpa variety) and three (Dia­
mante varieties) days when the floral buds were covered with pollen.
3.5.6. Autopolyploid
The use of autopolyploids has been implemented to increase the
yield of tomatillo crops. Autopolyploid consist in an acquisition of more
than two chromosomes sets by a genome duplication. The inducible
biological state in tomatillo is characterized by increasing effective
population size and allowing them to adapt and persist through het­
erogeneous landscapes (Ramírez-Godina et al., 2013).
3.5.7. Tissue culture
Khan and Bakht (2015) described the protocols for in vitro P. ixocarpa
micropropagation. Shoots of nodes of explants were planted into
Murashige-Skoog (MS) medium containing different concentration of
6-benzylaminopurine (0.5, 1.0 and 1.5 mg/L). Sprouting of the shoots
started between 7 and 8 days of inoculation. The 1.5 mg 6-benzylamino­
purine/L solution induced the maximum shoot length (3.45 cm).
Contreras and Almeida-Puentes (2003) analyzed the micro­
propagation of explants (hypocotyls) of P. ixocarpa L. The explants were
cultured on MS medium with inositol, nicotinic acid, glycine, pyridox­
ine, thiamine-HCl and sucrose. Eight days after incubation, small
white-greenish cells emerged from each explant. The propagation
continued and rooting was induced on the same medium. Finally, the
plants generate fruits (9–14/plant) and fertile seeds. García-Osuna et al.
(2015) stablished a protocol for the germination and micropropagation
of tetraploid tomatillo explants. The seeds germination was done
applying regulators (gibberellic acid) and organic acids (benzoic, sali­
cylic and sulfosalicylic acids), at concentration of 10− 2, 10− 4 and 10− 6
M. After 15 days in the MS basal medium, explants of the germinated
seedlings were sectioned. Hypocotyls, cotyledons and nodal segments
were used as explants. The explants micropropagation was performed in
MS medium (100 g myoinositol/L, 1 mg thiamine-HCl/L, 1 mg
pyridoxine-HCl/L, 40 mg adenine/L, 30 g sucrose/L, 4 g phytagel/L) and
combined with growth regulators (benzyladenine, kinetin and naph­
thaleneacetic acid). The best treatment was 3 mg benzyladenine pro­
ducing 9.5 bubs/explant.
On the other hand, leaf discs or hypocotyl and epicotyl explants of
P. ixocarpa were plated in MS medium with sucrose (3%), Gelrite
(0.25%) or agar (0.8%), vitamins (B5) and 2,4-dichlorophenoxyacetic
acid (4 mg/L). Nodules formed on the explant were moved to basal
MS medium containing benzylaminopurine (1.0 mg/L) and indole-3-
acetic acid (0.5 mg/L). After 6–8 weeks of incubation (26 ◦ C), organo­
genesis or embryogenesis shoots formed on the nodule. Shoots formed
plants, which can be transferred to a peat-soil mix in the protected
cultivation. For micropropagation, single nodules were transferred into
vessels containing MS medium, vitamins (B5), Gelrite (0.25%), and su­
crose (10 g/L. These nodules rapidly developed sprouts from buds
J.E. González-Pérez and J.Á. Guerrero-Beltrán
(Moriconi et al., 1990).
3.6. Maturation of fruits
Benito-Bautista et al. (2015) analyzed the relationship between color
and maturity characteristics of different P. ixocarpa varieties after the
fruit harvest. The varieties analyzed were San Martin, Tecozautla,
Rendidora and Diamante cultivars. Fruits were harvested after five
weeks; two weeks later, fruits of San Martin and Tecozautla varieties did
not have significant differences in color parameters. Fruits of Rendidora
and Diamante cultivars started to loss their green color due to an in­
crease of the a* color value (from − 14.1 to − 7.0), the green color was
turning pale. The Hue value was positively correlated with fruit texture
(less firmness).
4. Chemical composition and nutritional value
4.1. Minerals and vitamins
Table 4 shows some minerals found in fresh tomatillo fruits. Potas­
sium (K) is the main mineral existing in tomatillos. Fresh fruits also
contain Mg, Ca, Na, Fe, Zn, Mn, Cu and Se. Therefore, tomatillo is source
of K, P, and Mg. On the other hand, as a source of vitamins, fresh to­
matillo contains 440 μg thiamin, 0.035 mg riboflavin, 1.85 mg niacin,
0.15 mg pantothenic acid, 0.056 mg vitamin B6, 7 µg total folate, 0.38
mg α-tocopherol and 10.1 µg phylloquinone per 100 g of fruit
(Cárdenas-Castro et al., 2019; Shenstone et al., 2020). Therefore, as any
other fruits and vegetables, tomatillo may contribute with some
amounts of minerals and vitamins as nutrients to the humans’ daily
requirements.
4.2. Physicochemical and antioxidant characteristics
Physicochemical. Tomatillo fruits have around 92% of moisture
content and an average of 31–32 kcal/100 g fresh weight. The dry
matter of tomatillos contains 0.90–2.50% protein, 0.70–1.51% lipids,
0.8–1.07% ash and 0.40–1.90% total dietary fiber (Bock et al., 1995;
Cárdenas-Castro et al., 2019; McGorrin and Gimelfarb, 1998; Vdovenko
et al., 2018). Table 5 shows the physicochemical compositions of
different tomatillo varieties. The purple varieties have total soluble
solids (◦ Brix) content in the range 2.33–9.23%, pH in the range
4.05–4.56, and 0.8–1.1 g citric acid/100 g fruit. The purple fruits from
Jalisco, Mexico are less acid and sweeter than other commercial species
such as the one of Brazil, which has lower total phenolic components.
The green varieties have 5–6.89% total soluble solids and pH in the
range 3.79–4.29 (Table 2). Tomatillos are non-sweet fruits and some­
times very acid which make the adequate to be used for being used in
recipes for different dishes in Mexican and Guatemalan cuisines.
Antioxidants. As mentioned before, Table 5 shows concentration of
some chemicals in different tomatillo varieties. The concentration of
Table 4
Minerals in fresh tomatillo fruits.
Mineral Compositiona (mg/100 g)
K 268.0
P 39.0
Mg 20.0
Ca 7.0
Na 1.0
Fe 0.62
Zn 0.22
Mn 0.153
Cu 0.079
Se 0.0005
a
Shenstone et al. (2020).
8
Scientia Horticulturae 288 (2021) 110306
total anthocyanins, total phenolic compounds, flavonoids, tannins:
tomatidine in purple varieties is 0.0287–6.94 mg pelargonidin-3-
glucoside equivalents/g fresh weight, 0.15–10.08 mg gallic acid equiv­
alents (GAE)/g fresh weight, 0.1714–0.347 µg quercetin equivalents/g
dry extract, 0.122–1.49 mg catechin equivalents/100 g and
0.0249–0.0362 mg tomatidine/g, respectively (Shenstone et al., 2020).
The green varieties of tomatillo fruit contain 0.0253–1.5 mg
pelargonidin-3-glucoside equivalents/g fresh weight, 2.399–3.968 mg
GAE/g fresh weight, 0.338–1.46 µg quercetin equivalents/g dry extract,
0.126–1.884 mg catechin equivalents/100 g and 0.0223–3.81 mg
tomatidine/g, respectively. Khan et al. (2016) reported the concentra­
tion of polyphenol in n-butanol extracts of calyces, fruit, leaf and stem;
the polyphenol concentrations were 0.354, 20.11, 2.65 and 3.13 mg
GAE/g fresh weight, respectively. Cobaleda-Velasco et al. (2018)
determined the total flavonoids, total phenolic compounds, free radical
scavenging activity and iron reducing of different parts of P. ixocarpa
plant during its growing stage. The total flavonoids contents in stems
and leaves were 6.12–10.15 and 16.16–41.68 µg quercetin equiv­
alents/g dry extract, respectively. The total phenolic compounds in root,
stems and leaves were 0.37–0.55, 1.42–2.24 and 4.44–7.05 mg GAE/g
fresh weight, respectively. Free radical scavenging activity in root, stems
and leaves were 0.66–1.15, 0.45–0.90 and 0.09–0.25 mg 2,2-diphe­
nyl-1-picrylhydrazyl/mL, respectively. Total antioxidant capacities in
root, stems and leaves were 7.53–45.05, 26.23–68.71 and 55.82–216.31
µg ascorbic acid equivalent/mL, respectively. Thus, tomatillo consumes
in several ways can contribute to the antioxidants required in a human
diet.
4.3. Carotenoids
Fresh P. philadelphica and P. ixocarpa species of Puebla (Mexico) had
low concentration of β-carotene (2 and 21 µg/100 mg, respectively) and
lutein (12 and 1 µg/100 mg, respectively). Some studies have analyzed
the variation of this component during different cooking procedures of
tomatillos: boiling, roasting or steaming. β-carotene (a source of pro-
vitamin A) concentration did not show a significant variation due to
the boiling process. The microwave processing reduced twenty percent
the β-carotene in P. philadelphica. In the case of P. ixocarpa, microwaving
did not have effect in β-carotene nor lutein. Lutein concentration did not
show a significant difference due to cooking or boiling of P. ixocarpa and
P. philadelphica, respectively (Elizalde-González and Hernández-Ogarcí,
2007).
4.4. Terpenes and phenols
Tomatillos volatile compounds are methyl salicylate, β-caryophyl­
line, β-elemene, decanal, germacrene D, β-pinene, hexadecenoic acid,
methyl linolenate, benzyl alcohol and 3-carene (Gollapudi and Moto­
hashi, 2013). These terpenes and phenols, along with the acid taste and
other chemical compounds, give flavor (aroma, taste and sensation) to
tomatillos.
4.5. Steroidal lactones
Withanolides and physallins are lactones isolated from tomatillo.
The physallins reported in tomatillo are philadelphicalactone A, B, C and
D, 4β,7β,20R-trihydroxy-1-ocowitha-2–5‑dien-22,26-olide, 24,25-dihy­
drowithanolide D, 2,3-dihydro-3 βmethoxyixocarpalactone A, 2,3
dihydroixocarpalactone B, ixocarpanolide, ixocarpalactone A and B,
2,3-dihydro-3-methoxywithaphysacarpin, 18-hydroxywithanolide D,
and 2,3-dihydro-3β-methoxyixocarpalactone B, (Gollapudi and Moto­
hashi, 2013; Serdar and HanGamze, 2018). On the other hand, with­
anolides are a group of C-18 steroidal lactones; these compounds have
demonstrated anti-inflammatory, antimicrobial, anticancer, immuno­
regulatory, trypanocidal and leismanicidal activities (Huang et al.,
2019). Gu et al. (2003) isolated ixocarpalactone A from fresh fruit (143
 J.E. González-Pérez and J.Á. Guerrero-Beltrán
Table 5
Physicochemical and antioxidant characteristics of some genotypes of P. ixocarpa fruit.
Fruit color Location Genotypes Physicochemical attributes Total Total Phenolic Flavonoidsc Tanninsd Tomatidinee Reference
anthocyaninsa compoundsb

Purple Jalisco (Mexico) 17–196 ICTS-UDG 7.73◦ Brix, pH = 4.56, 1.07 g citric acid/ 6.01 4.68 González-Mendoza et al.
tomatillo 100 g (2011)
14–130 ICTS-UDG 5.07◦ Brix, pH = 4.39, 0.8 g citric acid/100 6.35 8.49
g
13–052 ICTS-UDG 7.87◦ Brix, pH = 4.39, 0.91 g citric acid/ 3 7.35
100 g
08–010 ICTS-UDG 7.93◦ Brix, pH = 4.10, 1.10 g citric acid/ 3.5 9.65
100 g
011–040 ICTS- 8.40◦ Brix, pH = 4.10, 0.97 g citric acid/ 1.5 7.89
UDG 100 g
012–048 ICTS- 9.23◦ Brix, pH = 4.11, 0.90 g citric acid/ 2.2 7.68
UDG 100 g
9–224- ICTS-UDG 3.9 10.08 González-Mendoza et al.
(2010)
9–032- ICTS-UDG 6.94 8.34
013–052 ICTS- 5 7.31
UDG
1–001- ICTS-UDG 5.4 5.5
2–002- ICTS-UDG 5.4 5.3
Guerrero L17 0.0292 2.094 0.347 1.49 0.0326 González-Chavira et al. (2019)
(Mexico)
Jalisco (Mexico) L41 0.0287 2.497 0.297 1.246 0.0249
9

Morelos (Mexico) L167 0.0318 2.519 0.305 0.339 0.0312

L171 0.048 3.244 0.1714 0.122 0.0304
Brazil Commercial 2.33◦ Brix, pH = 4.05, 0.91 g citric acid/ 0.1509 Curi et al. (2017)
100 g
Green Oaxaca (Mexico) Rendidora 5.81–6.45◦ Brix, pH = 3.79–4.15 1.5 1.46 Rosas-Medina et al. (2020)
tomatillo
Puebla (Mexico) L25 0.0326 3.606 0.338 0.331 0.0326 González-Chavira et al. (2019)
L30 0.044 2.924 0.351 0.297 0.0375
L37 0.0559 3.656 0.412 0.126 5.59
Jalisco (Mexico) L70 0.0517 3.832 0.3416 1.617 0.0306
L79 0.0292 2.399 0.3919 1.496 3.81
Morelos (Mexico) L86 0.0253 2.979 0.5117 0.345 0.03778
L89 0.0481 3.358 0.376 0.322 0.0223
L94 0.0271 2.6 0.3402 1.38 0.0365
L97 0.0459 3.206 0.4084 1.884 0.0226
L182 0.0722 3.968 0.6123 0.786 0.0374
a
mg pelargonidin-3-glucoside equivalents/g fresh weight.

Scientia Horticulturae 288 (2021) 110306

b
mg gallic acid equivalents (GAE)/g fresh weight.

c
µg quercetin equivalents/g dry extract.

d
mg catechin equivalents/g.

e
mg tomatidine/g; inhibition DPPH radical.
J.E. González-Pérez and J.Á. Guerrero-Beltrán
ppb). Su et al. (2002) investigated an ethyl acetate-withanolides extract
of leaves and stems of P. philadelphica. The extracts had phidelphica­
lactones A and B, ixocarpalactone A and B, withaphysacarpin,
18-hydroxywithanolide D, (2S,3S,4R,9E)− 1,3,4-trihydroxy-2- [(2′ R)−
2′ -hydroxytetracosanoylamino]− 9-octadecene, (2S,3S,4R)− 2-[(2′ R)−
2′ -hydroxytetracosanoylamino]− 1,3,4-octa-decanetriol, and (2S,3S,4R)−
2-tetracosanoylamino-1,3,4-octadecanetriol (chlorophyllide a). Choi
et al. (2006) obtained ethyl acetate extract and reported ixocarpalactone
A and B, philadelphicalactone B and withaphysacarpin. They also re­
ported that extracts were potent inducers of quinone reductase (cancer
chemoprotective activity for colon cancer cells). Ixocarpalactone A
induced apoptosis in SW480 cells, which suggest that tomatillo shows
potent antiproliferative and apoptotic activity in colon cancer cells.
4.6. Sucrose esters
Zhang et al. (2016) analyzed the chemical composition of sticky
material (a wax-like sticky material on pericarp) on tomatillo surface.
Chemical characterization of the sticky material yielded five sucrose
esters. 57.4% of sticky material was 2-O-decanoyl-3,10,30-tri-O-isobu­
tyryl sucrose and 2-O-decanoyl-3-O-(2-methylbutyryl)− 1′ ,3′ -di-O-­
isobutyryl sucrose. 6.8% of sticky material was 2,
3-di-O-decanoyl-1´-O-isobutyryl sucrose (C36H64O14) and 2,3-di-O-de­
canoyl-1´-O-(2-methylbutyryl) sucrose (C37H66O14). 4.8% of stick ma­
terial was 2,3-di-O-decanoyl sucrose. The five sucrose esters (100
µg/mL) from the sticky material and showed anti-inflammatory activity.
The anti-inflammatory activity was confirmed by in vitro cyclooxygenase
enzymes inhibitory assays. The cyclooxygenase enzyme inhibitory ac­
tivity was similar to drugs as aspirin (108 g/mL), ibuprofen (12 g/mL)
and naproxen (15 g/mL).
5. Flavor components
McGorrin and Gimelfarb (1998) identified over 50 volatile com­
pounds in tomatillo fruits. The green flavor of fresh tomatillo is provided
by aldehydes and alcohols such as (Z)− 3-hexenal, hexanal, (E,E)− 2,
4-decadienal, nonanal, 1-hexanol and (Z)− 3-hexen-1-ol. In addition,
others compound of tomatillo flavor include esters (hydroxy and aro­
matic), aldehydes (with 8–12 carbons), decanoic acid and terpenes. The
process of cooking tomatillo in water at 100 ◦ C for 2 min exhibited a
7-fold reduction of aldehyde levels and generation of valeryl hydrox­
y‑ester and cumic alcohol.
6. Uses
6.1. Fruit
In the United States, Canada, Mexico and some Central and South
American grocery stores, tomatillos are sold fresh or canned (Small,
2012). Tomatillo is an essential ingredient in the Mexican and Guate­
malan cuisines (Serdar and HanGamze, 2018). In the Mexican cuisine,
tomatillos are used without calyces as vegetables. It could be eaten raw
or cooked in different ways for preparing “salsas” and/or different
recopies. The cooking process enhances the zesty, tart flavor and softens
the thick pericarp. Typically, tomatillos are blended with onion, garlic,
cilantro and salt for making “salsas” (or green sauce) and “moles” (green
sauces that have several ingredients) with different amounts of chili.
Jams (with seed or seedless), jelly or curries are also prepared with to­
matillo (Curi et al., 2017; Pérez-Herrera et al., 2020; Shenstone et al.,
2020; Small, 2012). “Salsas”, sauce and dressings prepared with toma­
tillo are used to garnish dishes such as “tacos” (tortilla wraps filled with
meats), “enchiladas” (folded tortilla filled with chicken covered with
cooked green sauce), tostadas (crispy fried tortillas) or “tamales”
(cooked maize dough filled with green salsa and chicken or pork)
(Rengers and Gallagher, 2017). Some recopies of guacamole include
tomatillos. In addition, tomatillo could be stewed, fried, baked, cooked
10
Scientia Horticulturae 288 (2021) 110306
with small pieces of meat from different animals, as well as for making
soups, and dressings (Small, 2012).
For increasing their storage life in many parts of the world, tomatillos
are dehydrated and rehydrated before consumption. Dehydrated toma­
tillos have a great potential because this process may maintain their
nutritional and medicinal properties. They have a slight sweet and sour
taste (depending on the season), and do not oxidize during the drying
process. Recio-Colmenares et al. (2019) exposed slices of tomatillo (0.4
mm thick) to solar drying (1083–1036 W/m2 during 13.5 h) or
convective drying with an air speed of 2 m/s at controlled temperatures
(40, 50 and 60 ◦ C) for 800, 650 and 500 min, respectively.
On the other hand, P. philadelphica fruit has been used in the tradi­
tional medicine. The fruits are used to relieve fever, cough and amyg­
dalitis. The medicinal characteristics are related with withasteroids
(Maldonado et al., 2011).
6.2. Parts of tomatillo plants
Extracts from tomatillo plants have been used for stomach discom­
forts, for blood purification (antidote against local poison) or medicine
to treat sore throat (Khan and Bakht, 2015). The calyces and tomatillo
fruits are used in the treatment of diabetes (Hasnin-Aboelnaga, 2017;
Maldonado et al., 2011). Some parts of the tomatillo plant have been
used due to their new gastronomic taste or as a bio-scavenger of metals.
Husks of tomatillos are used to improve rheological properties of bread
dough and to enrich the rice flavor (Serdar and HanGamze, 2018).
On the other hand, husks and growing plants have being used in
contaminated soils (industrial areas) to scavenger heavy metals such as
Ni, Cd and Cu and in contaminated waters for removing iron and
manganese (responsible of metallic taste) (García-Mendieta et al.,
2012).
7. Concluding remarks and future perspectives
An overview about the tomatillo fruit, and plant, was included in this
review. There are many studies about techniques implemented to in­
crease the production of tomatillo and certain bioactive compounds
under controlled conditions in greenhouses and in the field as well as
studies reporting the benefits of compounds contained in tomatillos.
However, there are few reports that accurately indicate the recom­
mended amounts of their use in traditional medicine. The compounds
present in plants, calyxes and tomatillo fruit are of great commercial
importance. Tomatillo fruits and parts of the plant have a promising
future. Tomatillo could be commercialized around the world to be
consumed as a vegetable and be processed for making products by the
food industry, but also in the pharmaceutical industry, and for helping
the environment for removal of heavy metals in contaminated waters or
soils.
Declaration of Competing Interest
Authors declare no conflict of interest associated with this research.
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