Food Chemistry 354 (2021) 129571

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Food Chemistry
journal homepage: www.elsevier.com/locate/foodchem

Metabolomic analysis and physical attributes of ripe fruits from Mexican

Creole (Persea americana var. Drymifolia) and ’Hass’ avocados
Ana L. Ramos-Aguilar a, Juan Ornelas-Paz a, Luis M. Tapia-Vargas b, Alfonso A. Gardea-Bejar c,
Elhadi M. Yahia d, José de Jesús Ornelas-Paz a, *, Jaime D. Perez-Martinez e,
Claudio Rios-Velasco a, Pilar Escalante-Minakata f
a
Centro de Investigación en Alimentación y Desarrollo A.C.-Unidad Cuauhtémoc., Av. Río Conchos S/N, Parque Industrial, C.P. 31570, Cd. Cuauhtémoc, Chihuahua,
Mexico
b
Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias., Av. Latinoamericana No. 1101, Col. Revolución, CP. 60500, Uruapan, Michoacán, Mexico
c
Centro de Investigación en Alimentación y Desarrollo A.C.-Unidad Guaymas, Carretera al Varadero Nacional Km. 6.6, Col. Las Playitas, C.P. 85480, Guaymas,
Sonora, Mexico
d
Universidad Autónoma de Querétaro, Facultad de Ciencias Naturales., Avenida de las Ciencias S/N, C.P. 76230, Juriquilla, Querétaro, Mexico
e
Universidad Autónoma de San Luis Potosí, Facultad de Enfermería, Av. Niño Artillero No. 130, Zona Universitaria, C.P. 78210, San Luis Potosí, Mexico
f
Universidad de Colima, Laboratorio de Bioingeniería, Km. 9 carretera Coquimatlán-Colima, C.P. 28400, Coquimatlán, Colima, Mexico

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

Keywords: The physicochemical properties, including nutrient and bioactive compound compositions, in fruit of four creole
Mexican avocados avocados (CA) from Mexico were determined and compared with those of ’Hass’ fruit. ’Hass’ pulp and some CA
Native fruit pulps contained similar concentrations of lutein, chlorophyll a, β-sitosterol and α-tocopherol. CA pulp contained
Nutrients
3.91–9.55% more oil than ’Hass’. Oil from CA pulp contained 10.10–26.79% more oleic acid than ’Hass’ pulp.
Phytochemicals
Antioxidants
However, CA were small (CA = 81.40–137.15 g, ‘Hass’ = 188.59 g) and their pulp contents were low (CA =
39.83–84.82 g, ‘Hass’ = 144.14 g). CA peels were very thin, making these avocado peels edible but prone to
mechanical damage. CA peels also contained higher concentrations and greater diversity of anthocyanins and
glycosylated quercetin compounds than ’Hass’ peels. Some CA were particularly rich in mannoheptulose and
perseitol. Consumption of CA, including their peel, might result in higher intakes of some nutrients and bioactive
compounds compared with ’Hass’ avocados.

1. Introduction Fernández-Gutiérrez, & Carrasco-Pancorbo, 2014; López-Cobo et al.,

2016; Morais et al., 2017). ’Hass’ peel anthocyanin content has been
The quality of avocado fruit is determined by many attributes, determined only in a few studies (Ashton et al., 2006; Cox, McGhie,
including contents of nutrients and bioactive compounds (Salazar-López White, & Woolf, 2004). Existing data suggest that the sugar composition
et al., 2020; Yahia & Woolf, 2011). These attributes have been studied in ’Hass’ pulp is variable and influenced by origin, ripening stage, and
almost exclusively in ’Hass’ fruit due to its economic value. However, postharvest storage conditions (Blakey, Tesfay, Bertling, & Bower, 2012;
there are important knowledge gaps regarding the contents of certain Liu, Robinson, Madore, Witney, & Arpaia, 1999). Total carotenoid and
nutrients and bioactive compounds in ’Hass’ avocados. The contents of chlorophyll contents of ’Hass’ pulp have been studied extensively, but
oils, fatty acids, sugars and phenolic compounds have been determined the profiles of these compounds in ’Hass’ pulp and peel have been
extensively in the pulp of ’Hass’ but little is known about the contents of scarcely studied (Ashton et al., 2006; Lu et al., 2009). The chlorophyll
either nutrients or bioactive compounds in avocado peels, which are a concentration is important in avocados because they are an industrial
significant agroindustry by-product generated at high volumes with source of oil, and some studies have demonstrated that chlorophyll and
many potential uses (Hurtado-Fernández, Pacchiarotta, Mayboroda, other compounds determine the oxidation rate of oils during storage

* Corresponding author.
E-mail addresses: ana.ramos@estudiantes.ciad.mx (A.L. Ramos-Aguilar), juanopmx@gmail.com (J. Ornelas-Paz), tapia.luismario@inifap.gob.mx (L.M. Tapia-
Vargas), gardea@ciad.mx (A.A. Gardea-Bejar), yahia@uaq.mx (E.M. Yahia), jornelas@ciad.mx (J.J. Ornelas-Paz), jdavidperez@uaslp.mx (J.D. Perez-Martinez),
claudio.rios@ciad.mx (C. Rios-Velasco), minakata@ucol.mx (P. Escalante-Minakata).

https://doi.org/10.1016/j.foodchem.2021.129571
Received 17 April 2020; Received in revised form 1 March 2021; Accepted 6 March 2021
Available online 13 March 2021
0308-8146/© 2021 Elsevier Ltd. All rights reserved.
A.L. Ramos-Aguilar et al. Food Chemistry 354 (2021) 129571

(Yang et al., 2013). Chlorophyll and chlorophyll derivatives also exert 2. Materials and methods
beneficial effects on human health (Hayes & Ferruzzi, 2020). There are
also knowledge gaps about the contents of phytosterols, tocopherols and 2.1. Chemicals and plant materials
other compounds in ’Hass’ tissues (Campos et al., 2020; Chun, Lee, Ye,
Exler, & Eitenmiller, 2006), although the contents of organic acids in Solvents (analytical and HPLC grade), reagents, and standards of
avocado pulp have received interest in recent years (Campos et al., phenolic compounds, sugars, organic acids, carotenoids, chlorophylls,
2020). Knowledge about the chemical compositions of avocado might be tocopherols, phytosterols, and fatty acids were purchased from Sigma-
useful for understanding the relevance of this food in human health and Aldrich (St. Louis, USA). Perseitol and volemitol were purchased from
nutrition as well as potential valorization of by-products from the avo­ Carbosynth Ltd (Newbury, UK).
cado processing industry. Thirty mature-green fruits of four CA types (Persea americana var.
Avocados exhibit great genetic diversity. More than 1000 avocado Drymifolia) were harvested from an experimental field in Uruapan,
varieties have been documented worldwide (University of California, Michoacan, Mexico. This is the largest avocado-producing area in the
2020). In Mexico, the largest producer and exporter of ’Hass’ avocados world. The fruits of each genotype were obtained from five trees, which
in the world, 23 avocado varieties have been registered, 624 different showed red buds, lanceolate leaves with anise-like aromas and green
avocado genotypes have been documented, and 58% of them belong to pedicels. The height of the trees was ~30 m and they were 50 years old.
the Mexican race (Persea americana var. Drymifolia) (Sánchez-Pérez, The CA fruit were named CA-1, CA-2, CA-3, and CA-4. Their genetic
1999). The number of avocado genotypes in Mexico is continuously identification will be reported elsewhere. Thirty ’Hass’ avocados were
increasing due to natural or induced crossing of avocado genotypes purchased at the mature-green stage from a local market. CA and ’Hass’
(Tapia-Vargas et al., 2017). Many of these Mexican genotypes are native fruit were ripened at ambient temperature (25 ◦ C). Fourteen fruits of
or creole avocados (CA). CA plants are highly resistant to diseases; each genotype were evaluated for firmness. Sixteen fruits were distrib­
therefore, they are mainly used as rootstock for the ’Hass’ cultivar uted in four individual groups of four fruits each. Each group was
(Tapia-Vargas, Vidales-Fernández, & Larios-Guzmán, 2017). CA trees considered a replicate. The fruits of each group were evaluated for
also require 25.00–52.63% less water than commercial avocado vari­ weight, length, diameter and external color. Then, the peel, pulp and
eties, which can require up to 2000 L of water per tree annually, seed of these fruits were separated and weighed to determine the yield of
depending on tree size and age (Sánchez-Pérez, 1999; Tapia-Vargas each tissue. The length and diameter of seeds were also determined. The
et al., 2017). The morphological properties, genetic variability, foliar pulp of fruits was pureed with a kitchen blender and evaluated for
characteristics and agronomic performance of CA trees have been tristimulus color. The peels were evaluated for thickness and then cut
extensively studied (Acosta, Almeyda, & Hernández, 2013; Rendón- finely. The pulp puree and peels were evaluated for dry matter (DM),
Anaya et al., 2019). However, CA fruit have been scarcely studied for titratable acidity (TA), pH, total soluble solids content (TSS), oil content,
their nutritional qualities although they are part of the Mexican diet. The fatty acids, sugars, organic acids, total and individual phenols, antioxi­
production of CA fruit has increased over the years and local consumers dant capacity (AC), carotenoids, chlorophylls, phytosterols, phytosta­
prefer CA over ’Hass’ fruit due to their superior organoleptic properties nols and tocopherols. These measurements were performed under dim
(Espinosa-Alonso, Paredes-López, Valdez-Morales, & Oomah, 2017). light. The values of TA, TSS and oil were reported in terms of fresh
These avocados are generally small, of variable shape and have a large weight (FW). The phytochemical content and AC were expressed on a
seed and a very thin peel (green, black or purple). This thin peel can be dry weight (DW) basis.
eaten along with the pulp, which likely increases the nutritional and
health-related value of CA, as demonstrated for peels of ’Hass’ fruit 2.2. Analysis of sugars and organic acids
(Wang, Bostic, & Gu, 2010). Unfortunately, the relevance of these
avocados as a source of nutrients has received little attention. Corrales- Aliquots of pulp puree (4 g) or peels (2 g) were homogenized with 25
García et al. (2019) demonstrated that the peel of CA contained more and 40 mL of water, respectively, using an Ultra Turrax IKA-T18 (IKA
anthocyanins than that of ’Hass’ avocados. The pulp or oil of some CA Works Inc., Wilmington, USA). The pulp extracts were defatted with
has higher concentrations of α-tocopherol and oleic acid and higher hexane (20 mL) by liquid–liquid extraction. The peel extracts were not
antioxidant and anti-inflammatory activities than those of ’Hass’ fruit defatted at this time. Then, the samples were sonicated for 2 min, and
(Espinosa-Alonso et al., 2017; Peraza-Magallanes et al., 2017). Oil from the aqueous phase was recovered by centrifugation (20,000×g, 10 min,
some CA also showed better sensory properties than that of ’Hass’ fruit 4 ◦ C). The aqueous extract was filtered through Whatman paper No. 2
(Castañeda-Antonio et al., 2015). Other quality attributes of CA remain and subjected to two steps of defatting using hexane (20 mL) as
unknown; however, it is possible to hypothesize that CA might show described above. Finally, the aqueous extracts were filtered using a
better quality traits, including higher concentrations of nutrients and nylon membrane with a pore size of 0.20 μm and injected (20 μL) into
bioactive compounds than ‘Hass’ fruit, as observed for wild/creole ge­ the HPLC system for identification and quantification of sugars and
notypes of other fruits and vegetables (e.g., cherries, mushrooms, ap­ organic acids.
ples, cactus stems, etc.) and their commercial counterparts (Sun, The analysis of sugars was performed using a Varian HPLC system
Janisiewicz, Nichols, Jurick, & Chen, 2017; Veberic, Slatnar, Bizjak, (Varian Inc., Walnut Creek, USA) equipped with a refractive index de­
Stampar, & Mikulic-Petkovsek, 2015). Creole and commercial geno­ tector (Star Model 9040). The sugars were separated in a Shodex SUGAR
types biosynthesize different levels and types of bioactive compounds SC 1821 (8.00 × 300 mm) (Showa Denko K.K., Tokyo, Japan) ion-
under stress as defense mechanisms; however, CA are more resistant to exchange column at 70 ◦ C. The mobile phase was HPLC-grade water
biological and physical factors, suggesting that they contain higher at a flow rate of 0.60 mL/min. The analysis of organic acids was per­
levels and greater diversity of bioactive compounds (Ahanger, Tomar, formed using an Agilent 1200 series HPLC system (Agilent Inc., Palo
Tittal, Argal, & Agarwal, 2017; Boscaiu, Donat, Llinares, & Vicente, Alto, USA) equipped with a diode array detector (DAD). The separation
2012). Unfortunately, CA fruit has not been systematically compared to was carried out in an Aminex HPX-87H (300 × 7.80 mm) ion-exchange
the omnipresent ‘Hass’ fruit. Thus, the objective of this study was to column (Bio-Rad Laboratories, Hercules, USA). The mobile phase was
compare the contents of nutrients and bioactive compounds and other composed of 5 mM H2SO4 and acetonitrile (90:10, v/v) and the flow rate
quality traits in some CA and ’Hass’ avocados. was 0.40 mL/min. Ascorbic acid was monitored at 260 nm, and the other
acids were monitored at 210 nm. Identification and quantification of
sugars and organic acids were performed using standard compounds.

2
A.L. Ramos-Aguilar et al. Food Chemistry 354 (2021) 129571

2.3. Phenolic compounds and antioxidant capacity μA and 3000, 200, and 60 V, respectively. The carotenoid- and
chlorophyll-rich extracts were also evaluated for tocopherols. α- and
Samples of pulp puree (4 g) or peels (2 g) were homogenized in δ-Tocopherols were isocratically separated in the same column using a
methanol (40 mL) containing 0.10% trifluoroacetic acid (TFA). The mixture of methanol:MTBE (9:1, v/v) as the mobile phase at 0.75 mL/
extracts were sonicated for 2 min and then centrifuged (15,000×g, 20 min. Tocopherols were monitored by a fluorescence detector (λEX = 295
min, 4 ◦ C). The supernatant was recovered and evaporated at 40 ◦ C nm, λEM = 320 nm). Reference compounds were used for the identifi­
using a rotary evaporator. The residue was dissolved in 25 mL of water cation and quantification of these compounds.
containing 0.10% TFA. The extract was evaluated for the contents of
total (TP) and individual phenols and AC. For individual phenols, the 2.6. Analysis of phytosterols and phytostanols
extract was filtered through a nylon membrane with a pore size of 0.45
μm and injected (20 μL) into the Agilent HPLC system described above. The extracts used for carotenoids, chlorophylls and tocopherols were
The separation of phenols was carried out in a ZORBAX XDB-C18 col­ also analyzed for phytosterols and phytostanols using the Agilent 1200
umn (4.60 × 150 mm) (Agilent Inc., Santa Clara, USA) at 30 ◦ C. Phenolic HPLC system described above but connected to an evaporative light
compounds were monitored at 280, 320, 350 and 520 nm. The mobile scattering detector (ELSD) (Agilent 1260 Infinity). The separation of
phase was composed of 2.00% acetic acid (solvent A) and acetonitrile these compounds was performed in the ZORBAX XDB-C18 column
(solvent B), according to the gradient proposed by Ramos-Aguilar et al. (Agilent Inc., Palo Alto, USA) described above at 25 ◦ C. The mobile
(2017). Identification and quantification of compounds were performed phase (1 mL/min) was composed of acetonitrile (solvent A) and meth­
using standard compounds. anol (solvent B) according to the following gradient: 70.00% A from 0 to
TP was determined using the Folin Ciocalteu method at 750 nm using 26 min, 20.00% A at 36 min and 20.00% A from 36 to 50 min. Evapo­
a Bio-Rad microplate reader Model 680 (Bio-Rad, Tokio, Japan). The ration and nebulizer temperatures were 80 and 50 ◦ C, respectively. The
results were expressed as mg of gallic acid equivalents per kg (mg GAE/ drying gas was nitrogen (1.20 L/min). Standard compounds were used
kg DW). AC was measured at 490 nm, according to the method of Wang for identification and quantification.
et al. (2010). The results were expressed as mmol of Trolox equivalent
per kg of sample (mmol TE/kg DW). 2.7. Miscellaneous measurements

2.4. Oil analysis The length and diameter of fruits and seeds were measured using a
Vernier caliper, and peel thickness was determined using a micrometer
Oil was extracted from peels and pulps by the Soxhlet method using (Mitutoyo Corporation, Kawasaki, Japan). The firmness (14 fruits per
hexane as the solvent. The oil content was determined gravimetrically. avocado type) was determined by a TA-XT2 texture analyzer (Stable
Aliquots (50 mg) of oil were mixed with 1 mL of 0.5 N KOH, shaken and Micro Systems, Godalming, UK) equipped with a flat-head cylindrical
kept at 90 ◦ C for 10 min. After cooling, 1 mL of 14.00% BF3 in methanol probe (3 mm i.d.) at a speed of 0.18 mm/s. The tristimulus color (L*, a* y
was added and the sample was maintained at 90 ◦ C for 5 min to complete b*) was evaluated in four equidistant points of intact fruits (equatorial
the methyl esterification of fatty acids. The reaction mixture was cooled, arrangement) or petri dishes containing pulp puree. DM was gravimet­
mixed with 3 mL of hexane and shaken in a vortex for 1 min. The sample rically determined. Aqueous extracts (20 mL) from pulp puree (5 g) or
was heated at 90 ◦ C for 3 min, cooled and centrifuged (18,000×g, 10 peels (0.50 g) were evaluated for TA by titration with 0.01 N NaOH until
min, 4 ◦ C). An aliquot (1 µL) of the organic phase was injected into an HP a pH of 8.30 was reached. This extract was also evaluated for the content
gas chromatograph (5890 model) equipped with a flame ionization of TSS and pH using a digital refractometer (ATAGO Co. Ltd., Osaka,
detector (FID) and an HP 88 column (60 m × 0.25 mm; 0.20 μm film Japan) and a potentiometer (Denver Instruments, Arvada, USA),
thickness) (Hewlett Packard Company, Wilmington, USA). The injector respectively.
and detector temperatures were 250 ◦ C and 280 ◦ C, respectively. The
starting temperature in the oven was 100 ◦ C, increasing to 230 ◦ C at 2.8. Statistical analysis
2.50 ◦ C/min and, then, to 240 ◦ C at 0.50 ◦ C/min. Nitrogen was used as
the carrier gas (0.50 mL/min) and the split ratio was 25:1. The flow rates The experiment was completely randomized. The data were analyzed
of air and H2 in the FID were 300 and 30 mL/min, respectively. The by one-way ANOVA and the means were separated by the Tukey-Kramer
identification and quantification of fatty acid methyl esters were carried test (p < 0.05). All data analyses were performed using MINITAB 17
out using reference compounds. (Minitab Inc., State College, USA). All the measurements were carried
out at least in triplicate.
2.5. Analysis of carotenoids, chlorophyll and tocopherols
3. Results and discussion
Carotenoids, chlorophylls and tocopherols were extracted from pulp
(4 g) and peels (2 g) according to Ornelas-Paz, Yahia, and Gardea-Bejar 3.1. Biometrical characteristics
(2007). The carotenoids and chlorophylls were simultaneously sepa­
rated in a C30 column (150 × 4.60 mm) (YMC Inc., Milford, USA) at CA fruits were smaller in size and weight than ’Hass’ fruits (Table 1).
15 ◦ C. The mobile phase (0.75 mL/min) was composed of water (A), These attributes varied among CA genotypes. The phenotypic differ­
methanol (B) and methyl tert-butyl-ether (MTBE) (C), according to the ences among avocado genotypes have been attributed to genetics but
gradient proposed by Cervantes-Paz et al. (2012). Carotenoids were environmental factors may also be involved (Salazar-García, Medina-
monitored at 452 and 470 nm and chlorophyll was monitored at 470 and Carrillo, & Álvarez-Bravo, 2016). Our values of weight and length for
665 nm. The identity of carotenoids, chlorophyll and chlorophyll de­ the CA were similar to those (81.40–194.60 g and 5.00–10.90 cm) re­
rivatives was confirmed by HPLC-MS, using a 6210 time-of-flight (ToF) ported for other CA (Acosta et al., 2013). The weight of fruit parts fol­
mass spectrometer (Agilent Inc., Palo Alto, USA) equipped with an at­ lowed the order of pulp > seed > peel for CA and of pulp > peel > seed
mospheric pressure chemical ionization (APcI+) interface and Mass for ’Hass’ (Table 1). The large seed size of CA is considered a disad­
Hunter Manager software (A.02.01). The APcI+-MS system was operated vantage because it reduces the proportion of pulp. The peel thickness of
in positive ion mode. High-purity nitrogen (99.99%) was used as the ’Hass’ avocados was similar to that (1.19–1.74 mm) previously reported
nebulizing (20 psi) and drying gas (5 L/min). Other APcI+-MS param­ (Salazar-García et al., 2016). CA peels were 3 to 5 times thinner than
eters were as follows: gas and vaporizer temperatures of 325 and 350 ◦ C, those of ’Hass’ fruit. The peel thickness of CA-1 was exceptionally thin
respectively; corona, capillary, fragmentor, and skimmer voltages of 4 and similar to that of grape peels (0.24–0.33 mm) (Segade, Giacosa,

3
A.L. Ramos-Aguilar et al. Food Chemistry 354 (2021) 129571

Table 1 3.2. Firmness and color

Biometrical characteristic, firmness, dry matter, total soluble solids (TSS),
titratable acidity (TA), pH, oil and tristimulus color of Creole (CA) and ’Hass’ All of the tested fruit ripened properly as judged by their firmness,
avocados. which was below 5 N, the recommended minimum value of firmness for
CA-1 CA-2 CA-3 CA-4 ’Hass’ avocado consumption (Meyer & Terry, 2008). To the best of our
Fruit weight 104.53 137.15 81.40 ± 103.11 188.59 knowledge, this quality attribute has not been reported for CA. The a*
(g) ± 2.97c ± 3.82b 2.49d ± 4.49c ± 10.50a and b* color values of pulp indicated green-yellow coloration. The
Proportion Pulp 63.61 61.98 49.24 ± 47.95 76.43 ± ’Hass’ pulp showed an a* value lower than those of CA, indicating a
of (%) ± 1.43b ± 1.38b 0.84c ± 0.71c 1.72a greener tone. The pulps of CA-3 and CA-4 showed the highest and lowest
Seed 28.49 29.20 40.71 ± 42.89 12.50 ±
± 0.46b ± 0.79b 1.47a ± 0.99a 1.92c
b* values, respectively, causing noticeable differences in pulp yellow­
Peel 6.73 ± 5.96 ± 7.74 ± 7.07 ± 13.34 ± ness in these fruits. The peels were purple-black in color. The L* value of
0.44bc 0.34c 0.55b 0.47bc 0.34a ’Hass’ peel was higher than that of CA, indicating that the CA were
Fruit 5.55 ± 6.08 ± 5.12 ± 5.45 ± 6.86 ± darker than ’Hass’. The color values of ’Hass’ fruit were similar to those
diameter 0.06c 0.10b 0.05d 0.07c 0.21a
(Pulp: L* = 61.72, a* = − 1.48, b* = 31.02; Peel: L* = 23.85, a* = 1.25,
(cm)
Fruit length 6.22 ± 8.47 ± 6.67 ± 6.65 ± 9.80 ± b*=9.10) reported by Calderón-Oliver et al. (2016). The color of CA
(cm) 0.13c 0.15b 0.16c 0.12c 0.51a puree has been scarcely reported. Recently, Rodríguez-Campos,
Seed 3.57 ± 4.20 ± 3.73 ± 4.01 ± 3.21 ± Hernández-Carranza, Ávila-Sosa, Ruiz-López, and Ochoa-Velasco
diameter 0.05 cd 0.07a 0.05bc 0.07ab 0.20d (2020) determined the color (L* = 70.10, a* = − 21.70 and b* =
(cm)
52.10) of CA puree from Puebla, Mexico. To the best of our knowledge,
Seed length 3.96 ± 4.14 ± 3.84 ± 4.63 ± 3.84 ±
(cm) 0.08b 0.10b 0.05b 0.15a 0.02b there are no reports on the tristimulus color of CA peels. The variability
Peel 0.34 ± 0.39 ± 0.36 ± 0.50 ± 1.56 ± of color in the tested fruits suggests variability in the contents of ca­
thickness 0.01c 0.02c 0.01c 0.02b 0.06a rotenoids, anthocyanins, chlorophylls and chlorophyll derivatives,
(mm)
which confer the characteristic colors to the peel and pulp of these fruits
Fruit 1.32 ± 1.52 ± 1.65 ± 2.74 ± 2.68 ±
firmness 0.20b 0.14b 0.14b 0.18a 0.15a (Ashton et al., 2006).
(N)
Dry matter Pulp 34.17 30.02 33.04 ± 35.13 26.6 ± 3.3. Sugars
(%) ± ± 0.31c 0.58b ± 0.33a 0.1d
0.29ab
Peel 25.18 23.67 30.32 ± 29.17 31.13 ±
Overall, the content of TSS in pulp and peel was higher in CA than in
± 0.73b ± 0.32b 1.18a ± 0.72a 0.42a ’Hass’ fruits (Table 1). Six sugars were identified in the tested pulps and
TSS (%) Pulp 5.60 ± 5.19 ± 7.13 ± 6.79 ± 4.07 ± peels (Table 2), although other sugars previously reported in avocados
0.26b 0.43b 0.03a 0.10a 0.10c were not identified (ribose, stachyose and raffinose) (Blakey et al.,
Peel 16.09 16.31 13.57 ± 10.95 8.18 ±
2012). The contents of these sugars had not been characterized in CA.
± 0.02a ± 0.06a 1.30ab ± 0.01c
1.19bc The contents of these sugars in the pulp of CA and ’Hass’ avocados
TA (% citric Pulp 0.03 ± 0.03 ± 0.02 ± 0.02 ± 0.02 ± totaled 5.69–23.83 g/kg DW and 10.47 g/kg DW, respectively. On the
acid) 0.00a 0.00ab 0.00ab 0.00ab 0.00b other hand, the contents of these sugars in peels of CA and ’Hass’
Peel 0.06 ± 0.06 ± 0.05 ± 0.06 ± 0.02 ± avocados were 18.54–53.76 g/kg DW and 23.69 g/kg DW, respectively.
0.00a 0.00a 0.00a 0.00a 0.00b
pH Pulp 6.53 ± 6.71 ± 6.77 ± 7.19 ± 6.33 ±
Blakey et al. (2012) observed similar (6.20–8.90 g/kg DW) concentra­
0.14bc 0.10b 0.02b 0.04a 0.03c tions of sugars in the pulp of ripe ’Hass’ avocados from South Africa.
Peel 6.48 ± 6.85 ± 6.63 ± 6.84 ± 6.25 ± Fructose was the most abundant sugar in the pulp of CA-1, CA-2, CA-4
0.03b 0.02a 0.07b 0.03a 0.02c and in the peel of all CA, with the content of this sugar in these tissues
Oil (%) Pulp 20.57 22.07 17.79 ± 23.43 13.88 ±
being higher than that in ’Hass’ tissues, confirming that CA were sweeter
± 0.17b ± 0.61c ± 0.40a 0.59d
0.28ab than ’Hass’ fruit, as anecdotally reported by local consumers. Interest­
Peel 2.21 ± 3.50 ± 2.71 ± 2.65 ± 0.67 ± ingly, mannoheptulose (MH) was the most abundant sugar in the pulp of
0.18c 0.05a 0.05b 0.07b 0.04d CA-3, representing 48.93% of the quantified sugars. The ’Hass’, CA-1
Tristimulus and CA-4 pulps showed the lowest MH contents. However, CA-1 pulp
color had the highest perseitol content, which was up to 54.41% higher than
L* Pulp 55.82 55.04 61.72 ± 48.97 65.99 ± that of the other avocados. The perseitol content in the pulp of ’Hass’,
± 0.23c ± 0.61c 0.18b ± 0.54d 0.29a
CA-2, CA-3 and CA-4 fruits was similar. Glucose and fructose were the
Peel 26.91 28.14 29.66 ± 28.84 36.02 ±
± 0.44c ± 0.39b ± 0.37b 0.41a most abundant sugars in ’Hass’ pulp, and their concentrations were
0.53bc similar to those reported by Campos et al. (2020) for ’Hass’ fruit from
a* Pulp − 6.04 − 7.42 − 12.92 − 3.51 − 15.48 Peru; however, they observed that the most abundant sugar in pulp was
± 0.22b ± 1.2b ± 0.23c ± 0.28a ± 0.12d perseitol. This difference could be a consequence of differences in
Peel 6.50 ± 6.19 ± 6.05 ± 6.36 ± 2.74 ±
ripening rates between fruits because MH, perseitol and sucrose
0.14a 0.14a 0.14a 0.19a 0.31b
b* Pulp 26.89 30.94 37.37 ± 23.27 34.47 ± decrease during ripening, while glucose and fructose increase (Blakey
± 0.39d ± 0.3c 0.75a ± 0.38e 0.50b et al., 2012). In general, the composition of sugars in avocados is highly
Peel 1.36 ± 0.41 ± − 0.19 3.60 ± 0.24 ± dependent on many factors, including season, storage time and condi­
0.11b 0.15c ± 0.09d 0.21a 0.38 cd
tions, variety, and origin of fruit (Blakey et al., 2012; Kilaru et al., 2015).
Values represent the mean of several individual measurements ± the standard CA peels contained up to 183.18% more perseitol than ’Hass’ peel. MH
error. Values in the same row with different letters are significantly different (p was the most abundant sugar in ’Hass’ peel, with the content being
< 0.05). similar to that of CA-3 peel. Some beneficial effects have been attributed
to this sugar; unfortunately, the peel of ’Hass’ fruit cannot be consumed
Gerbi, & Rolle, 2011), making peel removal difficult from this avocado due to its thickness, hardness and bitterness. In general, the peels of all
type and favoring the coconsumption of pulp and peel. Local peoples avocados had higher sugar content than pulp. The contents of antioxi­
consume the peel and pulp of these kinds of fruit, suggesting that the dant sugars such as MH and perseitol in avocado peels, including peels of
peels do not compromise their organoleptic properties, unlike ’Hass’ ’Hass’ fruit, have rarely been reported (Liu et al., 1999). Thus, our data
avocados, which have much thicker peels that are not consumed. contribute significantly to the knowledge on the chemical composition

4
A.L. Ramos-Aguilar et al. Food Chemistry 354 (2021) 129571

Table 2 peel of ’Hass’ and CA-3 fruits, were rich in 7C sugars (MH, perseitol and
Sugars (g/kg DW) and organic acids (g/kg DW) in pulp and peel of Creole (CA) volemitol), which have been shown to exert some beneficial effects on
and ’Hass’ fruit. human health. MH and perseitol exert anticancer effects. MH favors
CA-1 CA-2 CA-3 CA-4 ’Hass’ normal levels of circulating glucose and helps in weight control (Ramos-
Sugars
Aguilar et al., 2019). Overall, CA showed an exceptionally high content
Fructose Pulp 2.81 ± 9.46 ± 8.97 ± 2.71 ± 3.88 ± of 7C sugars.
0.11b 0.97a 0.09a 0.33b 0.04b
Peel 12.18 42.17 31.23 ± 8.46 ± 1.89 ± 3.4. Titratable acidity, pH and organic acids
± ± 0.35b 0.86d 0.03e
0.45c 0.75a
Glucose Pulp 1.63 ± 1.27 ± 1.94 ± 1.26 ± 4.98 ± The TA and pH values (Table 1) were similar to those previously
0.07bc 0.02c 0.09b 0.13c 0.06a reported for avocados (Jacobo-Velázquez, Castellanos-Dohnal, Cabal­
Peel 7.05 ± 5.13 ± 6.73 ± 5.87 ± 5.47 ± lero-Mata, & Hernández-Brenes, 2013). This is the first study to report
0.28a 0.08c 0.33ab 0.52ab 0.13bc the content of up to six nonphenolic acids in tissues of avocado fruit
Sucrose Pulp 0.25 ± 0.34 ± 0.14 ± 0.18 ± 0.09 ±
0.02ab 0.04a 0.01c 0.00bc 0.00c
(Table 2). The organic acid composition has been scarcely investigated
Peel 1.00 ± 1.39 ± 0.60 ± 1.61 ± 0.21 ± in avocado fruit and few recent studies are available in this regard
0.03a 0.16a 0.07c 0.02a 0.00d (Campos et al., 2020). Unfortunately, some acids previously reported in
Mannoheptulose Pulp 0.54 ± 1.12 ± 11.66 ± 0.34 ± 0.34 ± avocado fruit, especially some phenolic acids, were not identified in the
0.02c 0.15b 0.02a 0.02c 0.01c
tested tissues (Hurtado-Fernández et al., 2014). The contents of organic
Peel 1.07 ± 1.71 ± 12.83 ± 1.02 ± 15.01 ±
0.04b 0.09b 1.07a 0.07b 0.15a acids in the pulp of CA and ’Hass’ fruits totaled 17.13–255.70 g/kg DW
Perseitol Pulp 2.04 ± 1.11 ± 1.02 ± 1.20 ± 0.93 ± and 61.22 g/kg DW, respectively. The total acid contents in peels of CA
0.05a 0.14b 0.04b 0.19b 0.01b and ’Hass’ avocados were 91.26–113.58 g/kg DW and 137.12 g/kg DW,
Peel 2.54 ± 3.03 ± 1.76 ± 1.58 ± 1.07 ± respectively. The composition of organic acids in CA had not been
0.11a 0.06a 0.30b 0.07b 0.01c
Volemitol Pulp ND* ND 0.10 ± ND 0.25 ±
determined previously. The concentrations of these acids were similar to
0.01b 0.01a those (succinic = 2.52–4.79 g/kg DW, quinic = 3.30–3.69 g/kg DW,
Peel ND 0.33 ± 0.44 ± ND 0.04 ± malic = 2.81–5.51 g/kg DW, citric = 3.38–4.40 g/kg DW) reported by
0.10a 0.05a 0.01b Campos et al. (2020). The contents of some organic acids in CA were
Organic acids significantly different from those of ’Hass’ avocados. Succinic and malic
Succinic Pulp 26.42 17.27 232.19 5.23 ± 44.10 ± acids were the most abundant acids in the tested avocados. Interestingly,
± ± ± 2.24a 0.32d 0.74b the content of succinic acid in CA pulps correlated well with that of oil
2.16c 3.81c
(R = − 0.85) and MH (R = 0.99). The oil and MH contents also correlated
Peel 68.04 11.24 287.69 17.21 123.75
± ± ± 7.96a ± ± 2.13b well with each other (R = − 0.84). For CA, fruit showing a high succinic
3.15c 0.88d 1.62d acid content had a low content of oil and a high content of MH. It has
Fumaric Pulp 0.43 ± 2.62 ± – 1.19 ± 1.01 ± been suggested that sugars are related to the accumulation of oil in
0.04c 0.09a 0.05b 0.02b
avocadoes during fruit growth (Liu et al., 1999). High expression levels
Peel 0.20 ± 0.82 ± 0.25 ± 0.68 ± 0.08 ±
0.01c 0.04a 0.02c 0.04b 0.01d of transketolase and transaldolase in plastids of avocado pulp contribute
Quinic Pulp 2.46 ± 1.05 ± 6.59 ± 1.63 ± 1.16 ± to the synthesis of MH, thus, C7 sugars such as MH could favor oil
0.14b 0.08c 0.32a 0.02c 0.06c accumulation (Ibarra-Laclette et al., 2015; Kilaru et al., 2015). The
Peel 9.78 ± 1.89 ± 20.71 ± 8.14 ± 5.38 ± relationship between the contents of succinic acid and oil suggests that
0.05b 0.00c 0.75a 0.57b 0.11c
the acid might be formed from β-oxidation, as inferred by others (Kilaru
Malic Pulp 2.68 ± 10.21 3.90 ± 7.51 ± 4.48 ±
0.14d ± 0.10c 0.01b 0.10c et al., 2015). The high content of succinic acid in the tested avocados
0.33c might contribute to the health-related properties attributed to this fruit
Peel 21.36 85.56 31.16 ± 60.02 5.48 ± such as anticonvulsant, hypocholesterolemic, hypoglycemic and anti­
0.79c 0.41e
± ± ±
diabetic properties (Venditti et al., 2016). Low concentrations of
1.15d 1.28a 0.03b
Citric Pulp 12.42 4.60 ± 12.53 ± 0.82 ± 10.14 ±
ascorbic acid were observed in pulp (0.07–0.17 g/kg FW). These con­
± 0.05c 0.25a 0.11d 0.09b centrations were similar to that (0.11 g/kg FW) previously reported for
0.19a ’Hass’ avocados (Yahia & Woolf, 2011). CA-1 and CA-3 pulps contained
Peel 21.62 14.07 24.49 ± 5.21 ± 2.38 ± up to 1.7 times more ascorbic acid than that of ’Hass’ fruit. The human
0.68a 0.04d 0.07e
health-related effects of ascorbic acid are well known and include the
± ±
0.74b 0.19c
Ascorbic Pulp 0.57 ± 0.30 ± 0.57 ± 0.33 ± 0.33 ± prevention of some degenerative diseases, such as heart disease and
0.07a 0.01b 0.03a 0.02b 0.01b some forms of cancer (Ramos-Aguilar et al., 2019).
Peel ND ND ND ND ND
Oxalic Pulp 0.17 ± 0.33 ± ND 0.42 ± ND
3.5. Total and individual phenols and antioxidant capacity
0.00b 0.03a 0.03a
Peel 1.12 ± 2.02 ± ND 1.72 ± ND
0.06b 0.17a 0.19ab The pulp and peel of ’Hass’ avocados contained more TP and higher
AC than those of CA (Table 3), agreeing with the well-known high
Values represent the mean of four individual measurements ± the standard
error. Values in the same row with different letters are significantly different (p
astringency of ’Hass’ peels that prevents the consumption of this tissue.
< 0.05).*ND, not detected. Our TP and AC values in the peel and pulp of ’Hass’ avocados (Table 3)
were similar to those previously reported for the same tissues (Wang
et al., 2010). These attributes had not been determined in CA. Sixteen
of avocados. In this study, sucrose and volemitol were the least abundant
phenolic compounds were identified in our samples (Table 3). The most
sugars in all the types of avocados. Liu et al. (1999) also observed that
abundant phenolic compound in the pulp of ’Hass’ avocados was
sucrose was the least abundant sugar in the pulp of ’Hass’ avocados,
chlorogenic acid, which was not detected in the pulp of CA. The content
probably because this sugar is used as a precursor of hexoses involved in
of chlorogenic acid was exceptionally high in the peel of ’Hass’ avocados
fruit respiration (Kilaru et al., 2015; Liu et al., 1999). The content of
compared with that in the peels of CA. Epicatechin was most abundant
volemitol in avocado pulp and peel had not been reported previously.
phenolic compound in the pulp of CA, but it was not observed in the pulp
This study demonstrates that the tested fruits, especially the pulp and
of ’Hass’ fruit, probably due to the ripening stage of the fruit. Hurtado-

5
A.L. Ramos-Aguilar et al. Food Chemistry 354 (2021) 129571

Table 3
Total phenolic content (TP), antioxidant capacity (AC) and individual phenols (mg/kg DW) in pulp and peel of Creole (CA) and ’Hass’ fruit.
CA-1 CA-2 CA-3 CA-4 ’Hass’
ab c bc c
TP (mg GAE/kg DW) Pulp 277.96 ± 36.32 146.50 ± 15.05 213.19 ± 43.53 140.78 ± 17.87 500.81 ± 27.62a
Peel 26145.54 ± 2939.88c 2697.54 ± 519.43d 30576.97 ± 1825.80b 4121.05 ± 58.75d 48116.36 ± 3734.98a
AC (mmol TE/kg DW) Pulp 0.21 ± 0.03bc 0.54 ± 0.06b 0.44 ± 0.02b 0.08 ± 0.04c 1.12 ± 0.02a
Peel 199.73 ± 15.41c 84.94 ± 3.56d 249.87 ± 14.47b 8.05 ± 0.52e 404.83 ± 11.53a
Chlorogenic acid Pulp ND* ND ND ND 55.29 ± 2.34
Peel 63.72 ± 1.09b 8.12 ± 0.94c 31.69 ± 2.17bc 15.79 ± 0.92c 3007.62 ± 21.41a
p-Coumaric acid Pulp ND 0.55 ± 0.04a 0.52 ± 0.01a ND ND
Peel 3.84 ± 0.02b ND 9.61 ± 1.38a 9.10 ± 0.50a ND
Cinnamic acid Pulp 1.74 ± 0.13a 1.84 ± 0.03a ND 1.94 ± 0.21a ND
Sinapic acid Pulp ND ND 1.58 ± 0.08b ND 34.64 ± 0.95a
Peel ND 44.62 ± 11.68 ND ND ND
Homovanillic acid Pulp 6.76 ± 0.90a ND ND ND 5.08 ± 0.24a
Tyrosol Pulp ND ND ND ND 18.89 ± 0.96
Vanillic acid Pulp 9.17 ± 0.45a ND 1.52 ± 0.10b ND ND
Peel 19.49 ± 1.06 a 7.76 ± 1.13b 10.58 ± 0.63b 8.45 ± 0.26b ND
Procyanidin B1 Pulp 14.26 ± 1.23bc 13.53 ± 2.05c 7.88 ± 1.88bc 21.53 ± 0.91b 42.69 ± 2.02a
Peel 54.46 ± 3.10a 29.33 ± 1.19bc 25.16 ± 3.41c 41.61 ± 2.75ab ND
Procyanidin B2 Pulp 71.21 ± 0.22 ND ND ND ND
Peel 289.04 ± 6.05c ND 659.01 ± 57.79b 356.41 ± 24.98c 5767.73 ± 9.25a
Epicatechin Pulp 134.65 ± 2.97d 403.70 ± 10.46b 508.94 ± 27.33a 263.41 ± 13.17c ND
Peel ND 707.24 ± 86.15b ND ND 7031.14 ± 491.29a
Catechin Pulp ND 8.37 ± 0.37a 8.26 ± 0.08a ND ND
Peel ND 35.50 ± 11.93b 174.19 ± 8.26a ND ND
Quercetin-β-glucuronide Peel 1509.14 ± 106.09a 49.72 ± 9.16c 1056.64 ± 101.66b 1123.47 ± 93.00ab ND
Quercetin-D-galatoside Peel ND ND 363.58 ± 7.71a 12.51 ± 1.24b ND
Quercetin-D-glucoside Peel 486.06 ± 25.10a 283.85 ± 64.53b 317.75 ± 25.63b 567.64 ± 45.50a ND
Cyanidin-glucoside Peel 685.96 ± 98.31b 160.03 ± 19.15c 2368.12 ± 497.69a 234.00 ± 20.09c 651.71 ± 27.98b
Pelargonidin-glucoside Peel 533.41 ± 55.98a 130.42 ± 38.84bc 497.52 ± 161.28b 55.85 ± 8.37c ND

Values represent the mean of four individual measurements ± the standard error. Values in the same row with different letters are significantly different (p < 0.05).
ND*, not detected.

Fernández et al. (2014) observed that the content of epicatechin 3 peel. A similar concentration of this anthocyanin in the peel of ’Hass’
decreased up to 89% in the pulp of ’Hass’ fruit during ripening. They fruit (230.00 mg/kg FW) was reported previously (Ashton et al., 2006).
also demonstrated that the composition of phenolic compounds was Interestingly, the peels of CA also contained pelargonidin-3-glucoside,
highly dependent on avocado variety. ’Hass’ pulp contained tyrosol. which had not been previously identified in avocado peels. This is the
Recently, López-Cobo et al. (2016) identified a glycoside of this com­ main pigment in ripe strawberries. Our values of pelargonidin-3-
pound (tyrosol-hexoside-pentoside) in ’Hass’ pulp (22.80 mg/kg DW). glucoside for CA peels (16.29–134.31 mg/kg FW) were similar to
Tyrosol is abundant in olive oil and wine and seems to be involved in the those reported for raspberries (30.00–31.00 mg/kg FW) and black
prevention of cardiovascular, metabolic and, neurodegenerative dis­ mulberries (116.00 mg/kg FW) (Veberic et al., 2015). The presence of
eases and some forms of cancer (Rodríguez-Morató et al., 2016). Only this anthocyanin in CA might explain the high a* values in peels of these
CA peels contained glycosylated-quercetin forms, with quercetin-3- fruits compared with that of ’Hass’ peels. Our data demonstrate that the
β-glucuronide being the most abundant in CA-1, CA-3 and CA-4. Quer­ peel of avocado represents an important source of glycosylated phenols,
cetin-3-glucoside was the most abundant in CA-2 peel. López-Cobo et al. especially some that are easily absorbed in the intestinal track.
(2016) identified other quercetin glycosides in ’Hass’ peel (quercetin-
diglucoside, quercetin-3-arabinosyl-glucoside and quercetin-3- 3.6. Oils and individual fatty acids
rutinoside). Glycosylated quercetin was not identified in the pulp of
the avocadoes we tested. Up to 17 procyanidins have been identified The pulp and peel of CA contained 3.91–9.54% and 1.54–2.83%
previously in avocado peels (López-Cobo et al., 2016; Wang et al., 2010). more oil than those of ’Hass’ fruit (Table 1). This might also explain the
Only procyanidin B1 and B2 were identified in the peel and pulp of the preference of local consumers for CA because the oil content is an
tested avocados. Overall, the peels showed greater diversity and higher important determinant of the sensory properties of foods, especially
abundance of phenolic compounds than pulps. Other phenolic com­ avocados. The oil content in the pulp of the avocados we tested was
pounds have been identified in the pulp of ripe avocados, which were similar to that previously reported for avocados. Espinosa-Alonso et al.
not identified in our study. Hurtado-Fernández et al. (2014) reported the (2017) reported oil contents of 19.65–26.77% and 18.28% in pulps of
acids 4-hydroxybenzoic, syringic and ferulic, while López-Cobo et al. CA and ’Hass’ fruit, respectively. Corrales-García et al. (2019) reported
(2016) reported some glycosylated phenols in avocados, including p- oil contents in the pulp of some CA and ’Hass’ fruit of 16.20–32.32% and
coumaric acid glucoside, p-coumaric acid rutinoside, ferulic acid 15.80%, respectively. The oil content in CA peels was similar to that
glucoside, and sinapic acid-C-hexoside, among others. The differences in reported for peels of other avocado genotypes (2.40%) (Morais et al.,
the profile of these compounds in avocados may be due to differences in 2017). Thirteen fatty acids were identified in the oil of the tested
the activity of phenylalanine ammonia lyase (PAL). It has been reported avocados (Table 4). Generally, a limited number of fatty acids (i.e., oleic,
that the concentrations of some phenolic compounds, such as chloro­ palmitoleic, linoleic and palmitic acid) is reported in oil from avocado
genic acid, decrease during fruit ripening due to a decrease in PAL ac­ tissues (Amado et al., 2019; Corrales-García et al., 2019; Morais et al.,
tivity and an increase in the overall catabolism of fruit (Villa-Rodriguez 2017). These fatty acids have already been identified in avocado oil, and
et al., 2020). their order of abundance agreed with that reported for oil from pulp of
The content of anthocyanins in avocado peels has scarcely been re­ CA, ’Hass’ and other avocado genotypes (Campos et al., 2020; Corrales-
ported (Ashton et al., 2006). Cyanidin-3-glucoside was the most abun­ García et al., 2019; Morais et al., 2017). The fatty acid composition in oil
dant anthocyanin in the peels. ’Hass’ peel only contained this from CA peel had not been reported previously.
anthocyanin and the concentration was 2.6 times lower than that of CA- Much of the human health-related properties of avocados have been

6
A.L. Ramos-Aguilar et al. Food Chemistry 354 (2021) 129571

Table 4
Fatty acid (mg/g of oil) composition in oil from pulp and peel of Creole (CA) and ’Hass’ fruit.
CA-1 CA-2 CA-3 CA-4 ’Hass’

Saturated
Capric Pulp ND* ND ND ND ND
Peel 1.38 ± 0.09a 1.15 ± 0.03b 1.19 ± 0.01b 1.14 ± 0.04b 1.48 ± 0.03a
Lauric Pulp ND ND ND ND ND
Peel 2.49 ± 0.15b 2.17 ± 0.01bc 2.28 ± 0.04bc 2.13 ± 0.05c 2.99 ± 0.14a
Mirystic Pulp 2.73 ± 0.04a 2.70 ± 0.04a 2.62 ± 0.06a 2.55 ± 0.02a 2.57 ± 0.05a
Peel 3.53 ± 0.14b 3.10 ± 0.09b 3.35 ± 0.03b 3.24 ± 0.16b 4.93 ± 0.25ª
Pentadecylic Pulp ND ND ND ND ND
Peel 1.17 ± 0.01ab 1.00 ± 0.02c 1.06 ± 0.00bc 1.02 ± 0.05bc 1.31 ± 0.07ª
Palmitic Pulp 130.62 ± 7.11a 129.42 ± 4.16a 115.67 ± 4.36a 137.09 ± 4.72a 128.96 ± 5.04a
Peel 133.60 ± 1.31a 132.20 ± 4.62a 122.63 ± 3.89ab 111.62 ± 5.68b 125.05 ± 5.10ab
Margaric Pulp 0.84 ± 0.04b 0.77 ± 0.01b 0.88 ± 0.04a 0.93 ± 0.02a 0.72 ± 0.02b
Peel 1.08 ± 0.04a 0.90 ± 0.02bc 0.97 ± 0.01ab 0.89 ± 0.04c 0.94 ± 0.03bc
Stearic Pulp 7.83 ± 0.28b 5.92 ± 0.07c 8.52 ± 0.28ab 9.16 ± 0.18a 4.39 ± 0.21d
Peel 11.89 ± 0.31a 7.68 ± 0.27bc 12.67 ± 0.39a 9.38 ± 0.51b 7.21 ± 0.37c

Monounsaturated
Palmitoleic Pulp 40.12 ± 2.03b 61.44 ± 3.21a 32.27 ± 1.18b 39.60 ± 1.81b 63.27 ± 2.27a
Peel 48.52 ± 0.10b 57.80 ± 2.18a 32.32 ± 1.28c 33.35 ± 1.74c 50.62 ± 2.11ab
Heptadecenoic Pulp 0.48 ± 0.09a 0.35 ± 0.03a 0.08 ± 0.01b 0.35 ± 0.06a 0.24 ± 0.05ab
Peel 0.23 ± 0.02b 0.42 ± 0.14ab 0.45 ± 0.10a 0.45 ± 0.04ab 0.52 ± 0.14a
Oleic Pulp 460.80 ± 21.62a 410.42 ± 13.17ab 473.02 ± 15.43a 470.20 ± 13.34a 373.06 ± 8.65b
Peel 458.60 ± 4.2a 388.51 ± 13.81b 448.35 ± 17.74a 347.61 ± 16.72bc 310.95 ± 10.07c

Poliunsaturated
Linolelaidic Pulp 0.90 ± 0.01a 0.92 ± 0.01ª 0.93 ± 0.02a 0.93 ± 0.01a 0.92 ± 0.01a
Peel 0.81 ± 0.01c 0.95 ± 0.01bc 0.95 ± 0.00bc 1.00 ± 0.05ab 1.17 ± 0.16a
Linolenic Pulp 7.83 ± 0.32b 8.90 ± 0.47b 10.37 ± 0.34a 10.49 ± 0.28a 8.58 ± 0.18b
Peel 17.33 ± 0.39b 13.70 ± 0.54c 20.97 ± 0.75a 16.80 ± 0.28b 16.34 ± 0.57b
Linoleic Pulp 87.43 ± 4.02ab 90.80 ± 3.64ab 99.39 ± 3.84a 92.98 ± 1.98ab 84.39 ± 2.63b
Peel 128.42 ± 0.91a 109.74 ± 3.54b 138.20 ± 4.74a 104.13 ± 1.99b 89.15 ± 2.40c

Values represent the mean of four individual measurements ± the standard error. Values in the same row with different letters are significantly different (p < 0.05).
ND*, not detected.

related to their content of unsaturated fatty acids, especially to the ’Hass’=13.39 mg/kg FW) were higher than that (8.00 mg/kg FW) in the
concentrations of oleic, palmitoleic and linoleic acids. The oil from CA- peel of ’Hass’ avocados from New Zealand (Ashton et al., 2006). Studies
1, CA-2 and CA-3 pulps contained up to 26.79% more oleic acid than have shown that the consumption of avocados effectively increases the
that of ’Hass’ pulp. The oil from CA-1 and CA-2 peels contained up to level of lutein in serum, improving eye and cognitive health (Ramos-
47.48% more oleic acid than ’Hass’ peels. In our study, the oleic acid Aguilar et al., 2019). The pulp and peel of the tested avocados contained
content in ’Hass’ peel was similar to that (822.70 mg/100 g DW) in oil small concentrations of neoxanthin, violaxanthin and zeaxanthin. The
from peels of Brazilian avocados (Morais et al., 2017). The oil from the low concentrations of these carotenoids might be a consequence of their
pulp and peel of CA-2 and ’Hass’ fruits showed the highest concentra­ role as precursors for abscisic acid biosynthesis during ripening (Cher­
tions of palmitoleic acid. The oil from CA-3 tissues showed higher nys & Zeevaart, 2000; Ibarra-Laclette et al., 2015). Very low concen­
contents of linoleic acid than those of ’Hass’ fruit. Oleic acid is involved trations of α- (0.14–0.52 mg/kg DW) and β-carotene (0.09–0.14 mg/kg
in protein G-mediated signaling, causing a reduction in blood pressure DW) were also observed in the pulp of the tested avocados, similar to
(Teres et al., 2008). Palmitoleic acid has been related to the control of that reported in ’Hass’ pulp (Ashton et al., 2006; Villa-Rodriguez et al.,
hyperglycemia, hypertriglyceridemia and insulin sensitivity (Campos 2020). In contrast to pulps, the peels contained high concentrations of α-
et al., 2020; Ramos-Aguilar et al., 2019). Acetogenins, which are fatty and β-carotene. ’Hass’ peel showed the highest concentration of
acid derivatives, were not considered in this study (Ramos-Aguilar et al., α-carotene, which was up to 10.1 times higher than that in CA peels.
2019). However, the peel of ’Hass’ fruit is not edible and therefore does not
contribute to carotenoid intake. On the other hand, CA peels are edible
3.7. Carotenoids, chlorophylls, tocopherols, phytosterols and phytostanols and contribute to carotenoid intake. The low concentrations of these
carotenes might be related to the ripening process because the content of
The carotenoids lutein, neoxanthin, lutein-5,6-epoxide, cis-viola­ β-carotene in the pulp and peel of avocados decreased significantly
xanthin, 15-cis-zeaxanthin, zeaxanthin, 13-cis-lutein, 15-cis-lutein, and during ripening (Villa-Rodriguez et al., 2020). α- and β-Carotene are
α- and β-carotene were detected and quantified in the tested avocados precursors of vitamin A and exert many beneficial effects on human
(Table 5). Lu et al. (2009) observed low concentrations of neochrome, health (Ramos-Aguilar et al., 2019).
chrysanthemaxanthin and β-cryptoxanthin in the pulp of ’Hass’ Chlorophyll and its derivatives are important pigments in avocados;
avocados, which were not observed in our study. To the best of our however, few studies have reported their contents (Ashton et al., 2006;
knowledge, only the total carotenoid content has been reported previ­ Wang et al., 2010). Chlorophyll a was more abundant than chlorophyll b
ously in the pulp of CA (Méndez-Zúñiga et al., 2019). This is the first in the peels; however, the content of these pigments was similar in both
study reporting the content of individual carotenoids in the pulp and tissues (Table 5). Chlorophyll a and b were more abundant in CA-4 tis­
peel of CA. Lutein was the most abundant carotenoid in the pulp and sues than in the other avocados. Pheophorbide a and pheophytin a were
peel of the tested avocados, with the concentrations of this carotenoid observed in the pulps and peels, while chlorophyll a’ and pheophorbide
(1.75–3.97 mg/kg FW) being similar to that (2.20–2.93 mg/kg FW) b were only identified in peels. Ashton et al. (2006) observed pheo­
reported previously for ’Hass’ pulp (Ashton et al., 2006). The pulp of CA- phytins a and b and chlorophyllides a and b in avocado fruit. The
3 contained 41.10% more lutein than that of ’Hass’. The concentrations ripening-related degradation of chlorophyll leads to the formation of
of this carotenoid in the tested peels (CA = 15.74–30.28 and pheophytins and chlorophyllides (Ashton et al., 2006). Ripe avocados

7
A.L. Ramos-Aguilar et al. Food Chemistry 354 (2021) 129571

Table 5
Carotenoid, chlorophyll and chlorophyll derivatives, tocopherols, phytosterols and phytostanols content (mg/kg DW) in pulp and peel of Creole (CA) and ’Hass’ fruit.
CA-1 CA-2 CA-3 CA-4 ’Hass’

Carotenoids
Lutein Pulp 7.03 ± 0.58bc 9.31 ± 0.61b 12.02 ± 0.96a 4.98 ± 0.07c 8.52 ± 0.14b
Peel 58.58 ± 5.94b 68.05 ± 11.53b 59.02 ± 5.36b 103.79 ± 5.70a 45.39 ± 2.22b
Neoxanthin Pulp 1.67 ± 0.41ab 2.24 ± 0.84a 1.62 ± 0.62ab 0.44 ± 0.06b ND*
Peel 10.28 ± 0.15b 13.13 ± 0.53a 12.22 ± 0.92a 4.65 ± 0.13c ND
Lutein-5,6-epoxide Pulp 0.05 ± 0.01c 0.10 ± 0.01ab 0.13 ± 0.00a 0.12 ± 0.02ab 0.09 ± 0.00ab
Peel 0.50 ± 0.07b 0.59 ± 0.11b 0.73 ± 0.06ab 0.85 ± 0.04a 0.49 ± 0.05b
9’-cis-neoxanthin Pulp 0.83 ± 0.18ab 1.31 ± 0.34a 0.84 ± 0.07ab 1.03 ± 0.17ab 0.23 ± 0.03b
Peel 11.00 ± 1.25ab 15.67 ± 3.40a 15.36 ± 0.15a 15.48 ± 1.88b 5.74 ± 1.06b
cis-violaxanthin Pulp 0.71 ± 0.20ab 1.31 ± 1.34a 0.84 ± 0.07 ab 1.03 ± 1.14a 0.23 ± 0.03b
Peel 10.61 ± 0.04a 11.34 ± 1.00ab 12.79 ± 0.34a 5.03 ± 0.28c ND
15-cis-zeaxanthin Pulp 0.22 ± 0.01d 0.69 ± 0.05a 0.32 ± 0.01 cd 0.39 ± 0.00bc 0.48 ± 0.02b
Peel 1.50 ± 0.43ab 1.01 ± 0.12ab 0.70 ± 0.18ab 1.91 ± 0.28a 0.45 ± 0.12b
Zeaxanthin Pulp 0.25 ± 0.01a 0.24 ± 0.02 a 0.35 ± 0.06a 0.30 ± 0.03a 0.30 ± 0.03a
Peel 1.22 ± 0.02b 1.04 ± 0.10b 1.62 ± 0.17b 3.17 ± 0.01a ND
13-cis-lutein Pulp 0.16 ± 0.01b 0.19 ± 0.02ab 0.22 ± 0.02ab 0.27 ± 0.04a 0.22 ± 0.02ab
Peel 1.00 ± 0.13b 1.15 ± 0.25b 1.29 ± 0.00ab 1.68 ± 0.05a 1.18 ± 0.06b
15-cis-lutein Pulp 1.08 ± 0.11c 1.94 ± 0.09a 1.66 ± 0.09ab 1.60 ± 0.18abc 1.37 ± 0.02bc
Peel 5.35 ± 0.11b 6.38 ± 0.28b 7.64 ± 0.40ab 10.03 ± 0.57a 5.38 ± 0.89b
β- carotene Peel 4.58 ± 0.70ab 3.04 ± 0.50ab 2.15 ± 0.02b 4.73 ± 0.25ab 6.40 ± 0.05a
α- carotene Peel 1.52 ± 0.16b 0.73 ± 0.05b 0.97 ± 0.09b 1.36 ± 0.15b 7.39 ± 1.28a

Chlorophyll and chlorophyll derivatives

Chlorophyll a Pulp 53.57 ± 3.93b 75.40 ± 1.44a 73.89 ± 7.01a 84.54 ± 5.92a 69.01 ± 2.39ab
Peel 751.71 ± 70.01b 616.84 ± 90.72bc 471.33 ± 14.63c 1051.24 ± 13.87a 722.99 ± 90.42ab
Chlorophyll b Pulp 56.33 ± 3.65bc 71.36 ± 4.12b 65.33 ± 0.39bc 95.55 ± 8.07a 43.68 ± 1.41c
Peel 431.11 ± 39.12ab 390.34 ± 25.48bc 286.40 ± 9.17 cd 525.37 ± 23.34a 201.33 ± 1.54d
Chlorophyll a’ Peel 2.09 ± 0.18b 2.62 ± 0.67b 2.21 ± 0.16b 5.11 ± 0.52a 2.45 ± 0.21b
Pheophorbide b Peel 1.76 ± 0.26b 0.97 ± 0.07b 9.71 ± 2.19a 0.63 ± 0.16b 1.33 ± 0.18b
Pheophorbide a Pulp 0.13 ± 0.02a 0.10 ± 0.02a 0.16 ± 0.03a 0.21 ± 0.07a 0.23 ± 0.02a
Peel 48.68 ± 7.77bc 24.66 ± 5.24c 146.51 ± 7.21a 31.66 ± 9.29c 81.52 ± 7.87b
Pheophytin a Pulp 1.84 ± 0.00a 1.02 ± 0.35b 0.37 ± 0.04bc 0.17 ± 0.02c 0.57 ± 0.09bc
Peel 10.04 ± 2.27ab 11.96 ± 1.38bc 8.50 ± 0.19c 17.63 ± 0.97ab 24.78 ± 4.05a

Phytosterols
β-sitosterol Pulp 822.14 ± 15.82c 1248.26 ± 74.02a 1035.55 ± 11.96b 868.34 ± 49.56bc 897.33 ± 7.26bc
Peel 896.82 ± 17.22ab 859.12 ± 35.93b 605.07 ± 33.77c 1019.36 ± 43.07a 577.87 ± 5.96c
Stigmasterol Pulp 339.76 ± 18.88a 409.95 ± 18.06a 353.43 ± 11.31a 399.42 ± 13.22a 246.83 ± 5.54b
Peel 348.46 ± 8.91b 299.53 ± 26.56bc 283.22 ± 6.61c 415.82 ± 5.74a 115.83 ± 6.74d
Campesterol Pulp 139.01 ± 1.77ab 151.71 ± 7.63a 129.65 ± 2.60b 152.86 ± 4.71a 143.74 ± 2.45ab
Peel 239.56 ± 4.52a 217.53 ± 12.48ab 175.29 ± 11.48c 200.96 ± 1.31bc 165.70 ± 5.33c

Phytoestanols
Cycloartenol Pulp 508.98 ± 33.71c 801.47 ± 30.95a 510.56 ± 35.10c 306.80 ± 20.47d 639.33 ± 15.25b
Peel 356.11 ± 15.80a 293.11 ± 17.71b 175.18 ± 9.62c 208.34 ± 14.43c 183.95 ± 6.03c

Tocopherols
α Pulp 38.83 ± 3.50b 55.88 ± 2.46b 81.11 ± 7.01a 12.81 ± 0.58c 39.53 ± 0.77b
Peel 165.23 ± 11.01c 105.20 ± 7.59d 302.80 ± 13.21a 118.91 ± 4.72d 214.41 ± 4.53b
δ Pulp 1.53 ± 0.16a 1.33 ± 0.05ab 1.14 ± 0.07ab 1.00 ± 0.08b 1.52 ± 0.24ab
Peel 291.41 ± 22.70a 318.92 ± 12.53a 103.11 ± 11.02b 344.10 ± 25.34a 117.69 ± 1.81b

Values represent the mean of four individual measurements ± the standard error. Values in the same row with different letters are significantly different (p < 0.05).
ND*, not detected.

are rich in these compounds (Ibarra-Laclette et al., 2015). However, the CA-4 and the peel of CA-3 contained more stigmasterol than cyclo­
chlorophyll concentration in avocado peel decreases during ripening, artenol. Previous studies have already demonstrated that β-sitosterol
while the anthocyanin content increases (Ashton et al., 2006; Ramos- was the most abundant phytosterol in avocados (Campos et al., 2020).
Aguilar et al., 2019). Interestingly, the peel of CA-3, which showed The content of β-sitosterol in ’Hass’ pulp was similar to that in CA-1, CA-
the lowest chlorophyll content (757.73 mg/kg DW), also had the highest 3 and CA-4 pulp. The pulp of CA-2 contained up to 1.4 times more
anthocyanin concentration (2865.64 mg/kg DW). To our knowledge, β-sitosterol than that of ’Hass’ fruit. In general, the concentrations of
there is no information on the content of these compounds in CA. Our β-sitosterol in the tested fruits were similar (1451.00–1977.60 mg/kg
study demonstrated that some CA were rich in chlorophyll and chloro­ DW) to those reported for ’Hass’ pulp (Campos et al., 2020). The peels of
phyll derivatives, which have been shown to have antioxidant, anti­ CA-3 and ’Hass’ fruit showed less β-sitosterol than the peels of CA-1, CA-
cancer, antimutagenic and antigenotoxic properties (Ramos-Aguilar 2 and CA-4. ’Hass’ peel showed the lowest phytosterol content among
et al., 2019). the tested fruits. In most cases, phytosterols and phytostanols were
Four phytosterols were identified in avocado pulp and peel (Table 5). found in greater abundance in the pulp than in the peel. β-Sitosterol was
Other phytosterols have been identified in avocado oil (campestanol, clearly the most abundant phytosterol in avocado peels. The variation in
lanosterol, Δ5-avenasterol, Δ7-sitosterol, α-amyrin, lupeol + gramis­ the content of phytosterols among avocado genotypes is mediated by
terol, cycloeucalenol, 24-methylenecycloartanol, and citrostadienol) stress factors (e.g. dehydration, cold, etc.) and abscisic acid content,
(Berasategi, Barriuso, Ansorena, & Astiasarán, 2012). In general, the which induces the expression of sterol methyltransferases, key enzymes
order of phytosterol abundance in the tested fruits was β-sitosterol > for the biosynthesis of these compounds (Ibarra-Laclette et al., 2015;
cycloartenol > stigmasterol > campesterol. Only the pulp and peel of Valitova, Sulkarnayeva, & Minibayeva, 2016). This study is the first to

8
A.L. Ramos-Aguilar et al. Food Chemistry 354 (2021) 129571

report the phytosterol content in CA and avocado peels. The consump­ Chavez Millan, Laura V. Rodriguez-Gonzalez, and E. Ochoa for their
tion of phytosterol-rich foods has been related to the prevention of some technical assistance.
cancer forms and cardiovascular diseases (Ramos-Aguilar et al., 2019).
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