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Fatty acid and volatile organic compound profiling of avocado germplasm grown
under East-Central Florida conditions

Article  in  Scientia Horticulturae · February 2020

DOI: 10.1016/j.scienta.2019.109008

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Scientia Horticulturae
journal homepage: www.elsevier.com/locate/scihorti

Fatty acid and volatile organic compound profiling of avocado germplasm

grown under East-Central Florida conditions
Sajid Alia,b, Anne Plottoa, Brian T. Scullya, David Wooda, Ed Stovera, Nancy Owensa,
Cristina Pisania,c, Mark Ritenourd, Muhammad Akbar Anjumb, Aamir Nawazb, Safina Nazb,
Jinhe Baia,*
a
USDA, ARS, U.S. Horticultural Research Laboratory, 2001 S. Rock Road, Ft. Pierce, FL, 34945, USA
b
Department of Horticulture, Faculty of Agricultural Sciences and Technology, Bahauddin Zakariya University, Multan, 60800, Pakistan
c
USDA, ARS, Southeastern Fruit & Tree Nut Research Laboratory, 21 Dunbar Rd., Byron, GA, 31008, USA
d
Indian River Research and Education Center, University of Florida, 2199 S. Rock Road Ft. Pierce, FL, 34945, USA

A R T I C LE I N FO A B S T R A C T

Keywords: Worldwide avocado consumption is growing due to potential health benefits. Most research has been focused on
Persea americana ‘Hass’, which does not perform well in Florida. Fatty acids and volatile organic compounds (VOCs) were eval-
Volatile organic compounds uated in 14 avocado genotypes grown in East-Central Florida conditions and compared to ‘Hass’. Two saturated
VOC and five unsaturated fatty acids were detected. Total oil content (TOC) was 11–25%, with 58.2–71.5% un-
Fatty acid
saturated fatty acids (UFA). ‘FL Hass’ contained 20% TOC, near the average for genotypes tested, but with a low
Unsaturated fatty acid
UFA of 61.9%. ‘PA-6206’ (a ‘Hass’ x ‘Bacon’ seedling) and ‘35707’ (a ‘Catalina’ seedling) had higher TOC and
UFA than other genotypes, while the mostly West Indian types such as ‘Simmonds’, ‘Day’, ‘Pflume’ and ‘Miguel’
had low TOC. Detected VOCs were acetaldehyde, hexanal, (E)-2-hexenal, limonene, α-cubebene, α-copaene, and
β-caryophyllene. Most genotypes contained most of the VOCs. ‘Monroe’ lacked C6 aldehydes, and ‘Pflume’,
‘Bernecker-43’, ‘Lula’ and ‘35707’ lacked some or all sesquiterpenes.

1. Introduction
Due to increased consumer awareness of its healthy dietary value,
the demand for avocado (Persea americana Mill) has significantly in-
creased in the past few decades - more than 8% annually and 11-fold in
total from 1985 to 2017 in the U.S. (The Statistics Portal, 2018). The
fruit of avocado owes its nutritional value to high oil content and
abundance of unsaturated fatty acids (Meyer and Terry, 2008; Villa-
Rodríguez et al., 2011; Pedreschi et al., 2016). In comparison to
monounsaturated fatty acids (MUFA) which contain one unsaturated
carbon bond in the molecule, linoleic acid (C18:2) and linolenic acid
(C18:3) contain two and three unsaturated carbon bonds, respectively,
and are generally called polyunsaturated fatty acids (PUFA), with the
higher desaturation having more positive impact in human health and
chronic diseases (Benatti et al., 2004). Oil content and fatty acid dis-
tribution vary based on genotypes associated with the origin of the race
(Galvão et al., 2014; Pacetti et al., 2007; Yanty et al., 2011), growing
regions (Carvalho et al., 2015; Pedreschi et al., 2016; Donetti and Terry,
2014), harvest times (Ozdemir and Topuz, 2004), and postharvest
conditions (Meyer and Terry, 2010; Pedreschi et al., 2016; García-Rojas
et al., 2016).
Most avocado cultivars currently in production are renowned for the
rich nutty flavor and creamy texture (Obenland et al., 2012; Pisani,
2016). Although oil content, fatty acids combination and dry matter are
key attributes to fruit texture, fruit VOCs also play a critical role in
avocado flavor (Obenland et al., 2012; Galvao et al., 2016). The avo-
cado fruit generally has alcohols, lipid-derived aldehydes, alkanes, es-
ters, ketones, and terpene-based VOCs (Pereira et al., 2013). However,
the number, and concentration of VOCs may vary with cultivar, pro-
duction location, and analytical method of detection used (Galvao
et al., 2016; Obenland et al., 2012; Pereira et al., 2013).
Avocados can be grouped into three types or genetic races based on
their origin, which correlate with fruit morphology, cold tolerance (low
to high) and horticultural traits: West Indian (WI), Guatemalan (G) and
Mexican (M) associated with the subspecies P. americana var. amer-
icana, P. americana var. guatemalensis and P. americana var. drymifolia,
respectively (Nakasone and Paull, 1998). The concentration of fatty
acids varies widely among the races of avocado – from more than 20%

⁎
Corresponding author.
E-mail address: jinhe.bai@ars.usda.gov (J. Bai).

https://doi.org/10.1016/j.scienta.2019.109008
Received 17 July 2019; Received in revised form 4 November 2019; Accepted 5 November 2019
0304-4238/ Published by Elsevier B.V.

Please cite this article as: Sajid Ali, et al., Scientia Horticulturae, https://doi.org/10.1016/j.scienta.2019.109008
S. Ali, et al. Scientia Horticulturae xxx (xxxx) xxxx

for M, 8–15% for G and 3–8% for WI, but there are many inter-racial
hybrids, often called complex hybrids, that present intermediate char-
acteristics (Nakasone and Paull, 1998; Gómez López, 1998).
The most widely commercially grown cultivar is ‘Hass’, a G x M
hybrid, with an oil content ranging from 10 to 20% in Florida (Pisani,
2016). ‘Hass’ is widely produced in Mediterranean climates [California
(USA), Spain, Israel, Mexico, and South Africa] (Nakasone and Paull,
1998) but generally does not perform well in hot and humid tropical
climates such as Florida (Crane et al., 2016; Pisani, 2016). ‘Miguel’ (G-
WI), ‘Simmonds’ (WI), ‘Monroe’ (G-WI), ‘Lula’ (G-WI), and ‘Brogdon’
(complex hybrid) are the major commercial avocado cultivars in
Florida, along with over a dozen minor varieties (Tropical Research and
Education Center, 2008). The genotypes that we used in this research
included some of these cultivars and other cultivars and breeding lines,
which have shown promise in productivity and visual fruit quality
under the East-Central Florida lowlands and humid subtropical cli-
mates. The objective of this research was to explore and compare the
fatty acid and aroma flavor profiles of 14 avocados genotypes from
diverse backgrounds.
The test was repeated when fruit did not ripen after 7–10 days, i.e. if
fruit became shriveled, rubbery and never softened. A minimum of 20
fruits of uniform size were harvested in each genotype to obtain 20
fruits of similar ripeness for chemical analyses.
2.2. Fruit ripening and firmness determination
Whole fruit fresh weight was taken individually immediately after
harvest. Fruit were then exposed to 20 °C for up to 10 d for ripening.
Fruit firmness was measured with a TA-XT2® texture analyzer fitted
with a 5-cm flat-plate probe loaded with a 50-kg cell (Texture
Technologies Corp. and Stable Micro Systems Ltd., Hamilton, MA). The
probe was driven with a crosshead speed of 20 mm min-1, and force was
measured at 2.5 mm deformation. Measurements were taken every day
until firmness decreased to below 25 N, at which point fruit was con-
sidered “ripe” (Pisani et al., 2017). Internal firmness of the mesocarp
was then measured on a 15-mm slice cut equatorially in the middle of
each fruit using an 8-mm diameter convex probe driven to a 5-mm
depth in the tissue (Pisani et al., 2017).

2. Materials and methods
2.1. Plant material
Uniform sized, and defect-free fruit were harvested from 4 to 6 years
old mature avocado trees located at the USDA, ARS Picos Research
Farm in Fort Pierce, FL, USA (27°43′N, 80°41′W). A total of 14 avocado
genotypes namely ‘Bacon’, ‘Bernecker-43’, ‘Dade-3’, ‘Day’, ‘FL Hass’,
‘Lula’, ‘Miguel’, ‘Monroe’, ‘PA-6206’, ‘Pflume’, ‘Simmonds’, ‘Zutano’,
‘35706’, and ‘35707’ were used in the study (Table 1 and Fig. 1). ‘FL
Hass’, a G–M hybrid with fruit appearance and eating quality similar to
‘Hass’ was used as a reference along with published results on ‘Hass’
(Pisani et al., 2017).
Because maturity was unknown for these cultivars under East
Central Florida conditions, a series of preliminary ripening tests were
performed before the anticipated harvest date of each genotype. When
fruit had reached standard size for the genotype, four to five fruits were
harvested and placed at 20 °C. Fruit that softened uniformly over a
period of 7–10 days without external ripening treatment were deemed
mature and each genotype was harvested when it met these criteria.
2.3. Replicate setting, measurement of mesocarp dry matter, and sample
preparations for chemical analysis
For each genotype, 20 fruits were grouped into four biological re-
plicates, and each replicate consisted of 5 fruits. Because fruit did not
all reach the target softness at the same time, the four replicates of fruit
were sometimes sampled over several days. During ripening, whenever
5 fruits reached ≤ 25 N firmness, they were grouped in a replicate, and
the mesocarp tissue of 5 fruits was pooled, mixed and prepared for
chemical analysis. Once all four replicates were collected, extra fruit
were discarded.
Mesocarp tissue from each replicate was chopped into 5 mm cubes,
and sub-samples were prepared as follows:
For dry matter content measurement, 10 g of mesocarp cubes were
dried until constant dry weight in a vacuum oven at 55 °C.
For fatty acid analysis, a 500 mg homogenized mesocarp sample
(fresh weight) was transferred to a glass vial, and stored at −80 °C until
extraction and analysis for fatty acids, as described below.
For VOCs analysis, mesocarp slices and an equivalent amount (w/w)
of saturated NaCl solution were homogenized in a food blender, and 6 g

Table 1
Fruit weight, dry matter percentage and total oil contents of 14 avocado genotypes.
Genotypes Race1 Harvest time Fruit weight (g) Whole fruit firmness (N)14 Mesocarp firmness (N)15 Dry matter content (%) Total oil content (%)

Standard Hass G-M5 − 169–1952 – 4.4–6.7 3 22–37 4 11.0–20.4 4

Bacon G-M5 10/20/2016 382 ± 6.5 efg 16.03 ± 1.29 d 2.53 ± 0.50 bc 23.6 ± 1.0bcd 18.3 ± 1.2 cd
Bernecker-43 WI5-OP6 11/15/2016 432 ± 20.1 efg 23.17 ± 1.18 ab 2.17 ± 0.20 c 31.9 ± 1.9a 22.4 ± 0.9ab
Dade-313 WI5-OP6 9/20/2016 602 ± 20.0 c 6.08 ± 0.40 e 1.63 ± 0.04 c 25.1 ± 0.2bc 20.3 ± 0.7bc
Day G-WI7 9/9/2016 381 ± 8.0 fg 17.85 ± 1.49 cd 1.90 ± 0.26 c 19.0 ± 0.6ef 13.0 ± 1.19fg
FL Hass G-M7 10/25/2016 308 ± 8.6 g 18.42 ± 1.24 cd 3.28 ± 0.25 bc 26.2 ± 0.9b 20.4 ± 1.2bc
Lula G-WI8 10/25/2016 471 ± 1.9 def 16.42 ± 1.06 d 2.63 ± 0.17 bc 22.8 ± 1.7 cd 18.2 ± 1.5 cd
Miguel G-WI8 9/8/2016 479 ± 18.6 cdef 16.01 ± 1.36 d 2.05 ± 0.10 c 21.3 ± 2.1de 14.3 ± 0.9efg
Monroe G-WI8 12/13/2016 892 ± 32.7 a 25.36 ± 1.06 a 2.84 ± 0.26 bc 25.3 ± 0.7bc 21.5 ± 1.7bc
PA-62069 G-M 11/29/2016 358 ± 7.9 fg 19.69 ± 1.32 bcd 3.21 ± 0.34 bc 33.2 ± 1.2a 25.0 ± 0.8a
Pflume WI7 9/15/2016 874 ± 22.4 ab 16.51 ± 1.34 d 2.10 ± 0.15 c 18.2 ± 1.1ef 13.7 ± 0.6efg
Simmonds WI9 9/16/2016 762 ± 14.9 b 21.71 ± 0.78 abc 2.36 ± 0.16 bc 16.8 ± 1.2f 11.4 ± 0.8 g
Zutano G-M5 10/20/2016 380 ± 3.6 efg 18.96 ± 1.88 cd 2.20 ± 0.23 c 22.4 ± 0.6 cd 16.2 ± 0.8def
3570610 WI11-OP7 12/13/2016 574 ± 15.9 cd 18.82 ± 1.39 cd 4.06 ± 1.15 ab 22.8 ± 0.5 cd 16.7 ± 1.8de
3570710,12 WI11-OP7 12/13/2016 490 ± 8.9 cde 21.03 ± 1.33 bc 5.32 ± 1.13 a 25.3 ± 0.9bc 20.9 ± 1.4bc

Values are means ± standard errors of four replicates of 5 fruits each. Within each column, means followed by the same letter are not significantly different at
P ≤ 0.05 using the Duncan multiple range test. Values in Standard Hass were collected from published references and were not statistically compared to the
experiment data. OP = open pollinated.
1
Race: WI-West Indian; G-Guatemalan; M-Mexican; 2 Jorge et al. (2015) and Carvalho et al. (2015); 3 Arpaia et al., 2018; 4 Ozdemir and Topuz (2004) and Villa-
Rodríguez et al. (2011). 5Chen et al. (2009); 6Ploetz et al. (2001); 7open pollinated, only seed parent known; 8based only on visual observations; 9Described in Crane
et al. (2016); 10Hass x Bacon seedling; 11Catalina seedling; 12described in Ploetz et al. (2001), but name designations personal communication; and 13Dade see-
dling.14Whole fruit firmness measured by compression at the equator; 15Mesocarp firmness measured by penetration with a 8 mm probe on a 15 mm slice at the
equator.

2
S. Ali, et al.
Fig. 1. Color and shapes of 14 avocado genotypes used in the study. Color figure is available only in web version of the article.
of the homogenate was transferred into a 20-mL vial and crimp capped
with a silicone septum (Gerstel Inc., Linthicum, MD, USA). Samples
were stored at −80 °C until analyzed for VOCs by using gas chroma-
tography–mass spectrometry (GC–MS).
2.4. Fatty acids sample preparation and analysis
2.4.1. Lipid extraction and methylation
Frozen samples were thawed at room temperature and then heated
at 80 °C for 10 min. Fatty acids were extracted by using Pisani (2016)’s
methods. Briefly, lipids were extracted with a isopropanol:hexane 4:6
solvent mixture. After purification, fatty acids were methylated for
analysis by GC (Pisani, 2016).
2.4.2. Fatty acid analysis
Fatty acid methyl esters (FAMEs) were identified using the Supelco
37 component standard. The Supelco 37 component standard has 37
different FAMEs ranging from C6 to C24. The chromatographic re-
solution was maximized and used in setting up the GC method. Fatty
acid methyl ester (FAME) determination was performed using a GC
(Model 6890; Agilent Technologies, Santa Clara, CA) equipped with a
split/splitless injector in splitless mode, flame ionization detector (FID)
and a 100 m× 0.25 mm i.d. × 0.2 μm film thickness CP-Sil 88 for
FAME capillary column using helium carrier gas. The initial head
pressure of the carrier gas was 250 kPa and was electronically increased
throughout the run to maintain a constant column flow rate of the
carrier gas at 1.5 ml min−1. The oven temperature was held initially at
60 °C for 2 min, later increased at 16 °C min-1 to 170 °C and held for
6 min, and then increased to 250 °C at 4 °C min-1 for 10 min. The tem-
peratures of the injection port and the detector were 250 °C and 280 °C,
respectively. Fatty acid column eluents were positively identified by
matching their retention times with those of reference standards.
Calibration curve verifications were always within 15% of a known
concentration and the coefficient of determination showed that the
regression line perfectly fit the data (R2 > 0.95).The total oil content
(TOC) was the sum of all fatty acids on a fresh weight basis.
3
Scientia Horticulturae xxx (xxxx) xxxx
2.5. VOC sample preparation and analysis
Frozen sample vials were thawed under tap water, and loaded into
the autosampler (Model MPS2; Gerstel Inc.) equipped with a cooled
tray holder [a cooling plate (Laird Tech, Göteborg, Sweden) controlled
by a Peltier Thermostat (CTC Analytics AG, Zwingen, Switzerland)].
Samples were held at 4 °C in the cooled tray until analyzed. For ana-
lysis, samples were incubated for 30 min at 40 °C. A 2-cm solid phase
microextraction (SPME) fiber (50/30 μm DVB/Carboxen/PDMS;
Supelco, Bellefonte, PA, USA) was then exposed to the headspace for
30 min at 40 °C. After exposure, the SPME fiber was inserted into the
injector of a GC–MS (Model 6890 N/5975; Agilent) to desorb the ex-
tract for 15 min at 250 °C. The GC–MS equipment and settings were:
DB-5 column (60 m length, 0.25 mm i.d., 1.00 μm film thickness; J&W
Scientific, Folsom, CA, USA); the column oven was programmed to
increase at 4 °C min−1 from the initial 40 °–230 °C, then ramped at
100 °C min−1 to 260 °C and held for 11.7 min for a total run time of
60 min; helium was used as carrier gas at a flow rate of 1.5 mL min−1.
Inlet, ionizing source and transfer line were kept at 250, 230, and
280 °C, respectively; mass units were monitored from 30 to 250 m/z and
ionized at 70 eV. Data were collected using a data system (ChemStation
G1701 AA; Hewlett-Packard, Palo Alto, CA, USA). A mixture of C-5 to
C-18 n-alkanes was run at the beginning of each day to calculate re-
tention indices. VOCs were identified by comparing their mass spectra
with the authorized standard chemicals, the NIST mass spectral data-
base, and published retention indices (Linstrom and Mallard, 2018).
The standard curve for each VOC was based on spiking a serial dilution
of standard VOC mixture to the ‘Bernecker-43’ homogenate (as this was
the genotype with the least amount of VOCs), and concentrations were
calculated by using regression equations, obtained with five different
concentrations of each standard.
2.6. Statistical analysis
Analysis of variance was performed on fatty acids, fatty acid com-
position ratios, and VOCs and differences among genotypes were esti-
mated using Duncan’s new multiple range test. Differences were con-
sidered significant at the 5% significance level. Pearson correlation tests
S. Ali, et al. Scientia Horticulturae xxx (xxxx) xxxx

were performed between firmness (whole fruit and mesocarp), DM,
TOC and each fatty acids. Cluster analysis was carried out on fatty
acids, fatty acid composition ratios, and VOCs generated through
Euclidean distance, and a dendrogram was prepared using the un-
weighted group pair method for the arithmetic average. All statistical
analyses were performed using XLSTAT v. 2018.5 (Addinsoft, NY,
USA).
3. Results
3.1. Fruit weight, dry matter and total oil contents
Average fruit weight varied from 308 g (‘FL Hass’) to 892 g
(‘Monroe’) (Table 1 and Fig. 1). Whole fruit firmness values at sampling
time ranged from 6.08 N (Dade-3) to 25.36 N (Monroe), while mesocarp
firmness ranged from 1.63 N (Dade-3) to 5.32 N (‘35707’). Dry matter
was the lowest for ‘Simmonds’ (16.8%) and the highest for PA-6206
(33.2%). Similarly, total oil content was the lowest (11.4%) in ‘Sim-
monds’ and the highest (24.9%) in ‘PA-6206’ (Table 1). Those values
were generally within the range published for ‘Hass’ avocado.
3.2. Combination of fatty acids
Seven fatty acids were detected in the 14 avocado genotypes.
Among them, two were saturated [palmitic acid (C16:0), and arachidic
acid (C20:0)], and five were unsaturated [palmitoleic acid (C16:1),
oleic acid (C18:1), linoleic acid (C18:2), linolenic acid (C18:3), and
eicosenoic acid (C20:1)] fatty acids (Table 2). Oleic acid was the most
abundant fatty acid in 12 out of 14 genotypes, ranged from a minimum
of 28.08 % of TOC in ‘FL Hass’ to a maximum of 45.14 % in genotype
‘35707’ (Table 2). Palmitic acid was the second most abundant fatty
acid next to oleic acid in 12 out of 14 genotypes, and the most abundant
in the other two genotypes (‘FL Hass’, and ‘Bernecker-43’), with a range
from 22.94% in ‘35707’ to 31.56% in ‘FL Hass’ (Table 2). Linoleic acid
was ranked the third, and it ranged from 13.02–20.46 % in fruit of
genotypes ‘Monroe’, and ‘Simmonds’, respectively (Table 2). Palmito-
leic acid was lowest in genotype ‘35707’, and greatest in the fruit of ‘FL
Hass’ (Table 2). Linolenic acid and eicosenoic acid were detected only
in a few genotypes and at concentrations less than 1%: ‘Simmonds’,
‘35706’ and ‘35707’ produced linolenic acid, while ‘FL Hass’, ‘PA-6206’
and ‘35707’ produced eicosenoic acid (Table 2). The concentration of
arachidic acid was found lowest (2.61%) in ‘35706’, and highest
(7.84%) in ‘Zutano’ (Table 2).
3.3. Correlation among dry matter, total oil content and fatty acids
There was no significant correlation between whole fruit and me-
socarp firmness. On the other hand, there was a highly significant
correlation (r = 0.945; P < 0.0001) between DM and TOC (Table 3).
Among fatty acids, oleic was significantly and negatively correlated
with palmitic and palmitoleic acids (r = −0.783; P = 0.001, and r =
−0.859; P < 0.0001, respectively). Linolenic acid was positively cor-
related with linoleic acid (r = 0.587; P < 0.05) and arachidonic acid
was negatively correlated with linoleic and linolenic acids (r = −0.542
P = 0.045, and r = −0.595; P = 0.025, respectively) (Table 3).
3.4. Fatty acid composition ratios
Table 4 shows the proportions between unsaturated fatty acids
(UFA) and saturated fatty acids (SFA), and further dividing UFA to
monounsaturated fatty acid (MUFA) and polyunsaturated fatty acid
(PUFA). Among the UFA, oleic acid, linoleic acid, and palmitoleic acid
were the major fatty acids in the mesocarp tissue of avocado fruit. The
percentage of TOC represented by SFA ranged from 28.53 % (‘35707’)
to 41.84 % (‘Bernecker-43’); while, percent of UFA in the TOC ranged
from a minimum of 58.15% (‘Bernecker-43’) to a maximum of 71.47%
(‘35707’) (Table 4). The percentage of TOC represented by MUFA
ranged from 43.23–55.24% in the genotypes ‘Bernecker-43’, and ‘PA-
6206’, respectively. Similarly, the percentage of TOC represented by
PUFA also varied significantly ranging from 13.02% in ‘Monroe’ to
20.91% in ‘Simmonds’ (Table 4). All genotypes had > 1 MUFA/SFA
value, where the genotype ‘Bernecker-43’ had the lowest value (1.05),
while ‘PA-6206’ exhibited the highest (1.92) value. Similarly, PUFA/
SFA and (MUFA + PUFA)/SFA ratios were lowest in the genotype
‘Bernecker-43’ (0.36 and 1.41, respectively); while, ‘35707’ and ‘PA-
6206’ showed high values (Table 4).
3.5. Aroma VOCs
Seven VOCs including three aldehydes, one monoterpene, and three
sesquiterpenes were detected in the 14 avocado genotypes (Table 5). All
seven VOCs were detected in the genotypes ‘Day’ and ‘35706’, while
only four VOCs were detected in ‘Monroe’ and ‘Pflume’. ‘Day’, ‘Lula’
and ‘FL Hass’ had the highest total VOCs concentration, mostly from
acetaldehyde (‘Day’), hexanal (‘Day’ and ‘FL Hass’) and (E)-2-hexenal
(‘Lula’). ‘Bernecker-43’had the lowest concentration of total VOCs,
mostly explained by the lowest amount of acetaldehyde (Table 5). The
genotype ‘35706’ showed the lowest hexanal and (E)-2-hexenal. The

Table 2
Fatty acids content of mesocarp from 14 avocado genotypes on fresh weight basis.
Genotypes Palmitic acid Palmitoleic acid Oleic acid Linoleic acid Linolenic acid Arachidic acid Eicosenoic acid
[C16:0 (%)] [C16:1 (%)] [C18:1 (%)] [C18:2 (%)] [C18:3 (%)] [C20:0 (%)] [C20:1 (%)]

Standard ‘Hass’ 13.65–20.851-3 3.45–8.891-3 56.95–67.001,2 12.20–15.271-3 1.10–1.26 1-3 0.73–0.174, 5 0.37–0.484,5
Bacon 28.20 ± 3.17bc 9.37 ± 0.81ef 43.29 ± 0.35ab 13.27 ± 1.69 cd – 5.44 ± 1.04ab –
Bernecker-43 37.08 ± 2.87a 13.38 ± 0.48ab 29.84 ± 2.67fg 14.92 ± 0.40b-d – 4.76 ± 0.80bc –
Dade-3 32.21 ± 0.37ab 11.94 ± 0.29b-d 33.46 ± 0.71e-g 18.01 ± 0.97ab – 4.37 ± 0.08bc –
Day 27.00 ± 1.75bc 9.55 ± 0.40ef 41.75 ± 0.86a-c 17.20 ± 1.08a-c – 4.48 ± 1.50bc –
FL Hass 31.56 ± 1.63ab 15.34 ± 0.61a 28.08 ± 2.41 g 17.32 ± 0.89a-c – 6.51 ± 0.39ab 1.17 ± 0.82a
Lula 30.12 ± 1.46b 12.68 ± 1.36bc 34.63 ± 2.96d-g 16.09 ± 0.62b-d – 6.46 ± 0.13ab –
Miguel 28.20 ± 0.59bc 9.03 ± 0.52ef 39.23 ± 1.94a-e 18.61 ± 1.76ab – 4.90 ± 0.36bc –
Monroe 30.24 ± 3.04b 8.86 ± 0.50ef 42.48 ± 2.11ab 13.02 ± 0.31d – 5.39 ± 1.00ab –
PA-6206 22.84 ± 1.59c 10.00 ± 0.56d-f 44.42 ± 1.46a 15.79 ± 0.57b-d – 6.11 ± 0.14ab 0.81 ± 0.48ab
Pflume 28.71 ± 0.89bc 10.44 ± 0.48de 40.40 ± 2.25a-d 14.85 ± 1.23b-d – 5.58 ± 0.20ab –
Simmonds 31.66 ± 0.83ab 11.13 ± 0.44c-e 33.47 ± 2.73e-g 20.46 ± 2.39a 0.45 ± 0.45a 2.81 ± 1.11c –
Zutano 27.00 ± 1.13bc 14.02 ± 0.94ab 35.67 ± 2.08c-f 15.44 ± 0.40b-d – 7.84 ± 0.23a –
35706 32.37 ± 3.00ab 9.56 ± 0.90ef 36.89 ± 2.95b-e 18.18 ± 0.81ab 0.37 ± 0.38a 2.61 ± 1.17c –
35707 22.94 ± 1.06c 8.15 ± 0.70f 45.14 ± 1.74a 17.23 ± 1.29a-c 0.40 ± 0.41a 5.58 ± 0.30ab 0.53 ± 0.53ab

Within each column, means followed by the same letter are not significantly different at P ≤ 0.05 using the Duncan multiple range test. Values are means ± standard
errors of four replicates of five fruits each. Values in Standard Hass were collected from references and were not statistically compared with the experiment data. Data
from 1Pedreschi et al. (2016); 2Yanty et al. (2011); 3Meyer and Terry (2008); 4Ozdemir and Topuz (2004); 5Carvalho et al. (2015).

4
S. Ali, et al. Scientia Horticulturae xxx (xxxx) xxxx

Table 3
Pearson correlation coefficients among whole fruit firmness, mesocarp firmness, dry matter content, total oil content and different fatty acids of 14 avocado
genotypes.
Variables Whole Firm Pulp Firm DM TOC PA PTA OA LA LLA AA EA

C16:0 C16:1 C18:1 C18:2 C18:3 C20:0 C20:1

Whole Firm 1 0.385 0.173 0.128 0.001 −0.119 0.134 −0.261 0.278 0.018 0.125
Pulp Firm 1 0.253 0.364 −0.348 −0.306 0.307 0.070 0.615* −0.013 0.498
DM 1 0.945*** 0.034 0.184 −0.043 −0.376 −0.284 0.287 0.487
TOC 1 −0.027 0.114 0.049 −0.459 −0.254 0.350 0.511
PA 1 0.500 -0.783*** 0.039 −0.024 −0.397 −0.302
PTA 1 -0.859*** 0.032 −0.319 0.402 0.265
OA 1 −0.323 0.058 0.064 −0.074
LA 1 0.587* -0.542* 0.080
LLA 1 -0.595* −0.006
AA 1 0.359
EA 1

* = Significant at P ≤ 0.05, ** = Significant at P ≤ 0.01, *** = Significant at P ≤ 0.001. DM = dry matter, TOC = Total oil content, PA = Palmitic acid,
PTA = Palmitoleic acid, OA = Oleic acid, LA = Linoleic acid, LLA = Linolenic acid, AA = Arachidic acid, EA = Eicosenoic acid.

Table 4
Concentration of total saturated/unsaturated fatty acids and different fatty acid composition ratios of mesocarp from 14 avocado genotypes on fresh weight basis.
Genotypes Total SFA Total UFA (U, %) M/S P/S (M + P)/S
(S, %) (Ratio) (Ratio) (Ratio)
Total (%) Total MUFA (M, %) Total PUFA (P, %)

Bacon 33.64 ± 2.13b-d 66.35 ± 2.13b-d 52.63 ± 0.47a-c 13.72 ± 1.69 cd 1.58 ± 0.11b 0.42 ± 0.08c-e 2.00 ± 0.19a-c
Bernecker-43 41.84 ± 2.08a 58.15 ± 2.08e 43.23 ± 2.24 g 14.92 ± 0.40 b-d 1.05 ± 0.12d 0.36 ± 0.02e 1.41 ± 0.14f
Dade-3 36.58 ± 0.44bc 63.41 ± 0.44 cd 45.40 ± 0.63e-g 18.01 ± 0.97ab 1.24 ± 0.01 cd 0.49 ± 0.03a-d 1.73 ± 0.04de
Day 31.48 ± 0.50de 68.51 ± 0.50ab 51.30 ± 1.21a-d 17.20 ± 1.08a-c 1.63 ± 0.06ab 0.55 ± 0.03a-c 2.18 ± 0.09ab
FL Hass 38.07 ± 1.28ab 61.92 ± 1.28de 44.60 ± 2.01fg 17.32 ± 0.89a-c 1.18 ± 0.09 cd 0.46 ± 0.02b-e 1.63 ± 0.11ef
Lula 36.59 ± 1.40bc 63.40 ± 1.40 cd 47.31 ± 1.88c-g 16.09 ± 0.62b-d 1.30 ± 0.10b-d 0.44 ± 0.01b-e 1.74 ± 0.11de
Miguel 33.11 ± 0.36 cd 66.88 ± 0.36bc 48.27 ± 2.08b-g 18.61 ± 1.76ab 1.46 ± 0.08bc 0.56 ± 0.05ab 2.02 ± 0.13a-c
Monroe 35.63 ± 2.05b-d 64.36 ± 2.05b-d 51.34 ± 2.06a-d 13.02 ± 0.31d 1.46 ± 0.13bc 0.37 ± 0.02de 1.83 ± 0.15e-e
PA-6206 28.96 ± 1.50e 71.04 ± 1.50a 55.24 ± 1.36a 15.79 ± 0.57b-d 1.92 ± 0.13a 0.55 ± 0.04a-c 2.47 ± 0.17a
Pflume 34.29 ± 0.83b-d 65.70 ± 0.83b-d 50.85 ± 1.87a-e 14.85 ± 1.23b-d 1.49 ± 0.09bc 0.43 ± 0.03b-e 1.92 ± 0.12b-d
Simmonds 34.48 ± 0.30b-d 65.51 ± 0.30b-d 44.60 ± 2.30fg 20.91 ± 2.55a 1.29 ± 0.06b-d 0.61 ± 0.08a 1.90 ± 0.14b-d
Zutano 34.85 ± 1.32b-d 65.14 ± 1.32b-d 49.70 ± 1.21a-f 15.44 ± 0.40b-d 1.44 ± 0.09bc 0.45 ± 0.02b-e 1.89 ± 0.11b-e
35706 34.98 ± 2.02b-d 65.01 ± 2.02b-d 46.45 ± 2.08d-g 18.56 ± 0.68ab 1.35 ± 0.15b-d 0.54 ± 0.04a-c 1.89 ± 0.19b-e
35707 28.53 ± 0.82e 71.47 ± 0.82a 53.82 ± 2.12ab 17.64 ± 1.31a-c 1.90 ± 0.13a 0.62 ± 0.03a 2.52 ± 0.16a

Within each column, means followed by the same letter are not significantly different at P ≤ 0.05 using the Duncan multiple range test. Values are means ± standard
errors of four replicates of 5 fruits each. SFA (S) = Saturated fatty acids; UFA (U) = Unsaturated fatty acids; MUFA (M) = Monounsaturated fatty acids; PUFA (P) =
Polyunsaturated fatty acids.

Table 5
Concentration (μg kg-1) of different volatile organic compounds detected in 14 avocado genotypes.
Genotypes Acetaldehyde Hexanal (E)-2-Hexenal Limonene α-Cubebene α-Copaene β-Caryophyllene

1
Odor descriptor Solvent, fruity, fresh Fruity, green, stem-like Green, grassy, apple-like Lemon, orange Citrus, fruity Woody, spicy Spicy, woody
Retention indices 478 802 885 1048 1363 1400 1453
AT2(μg kg-1) 153 53 173 10-2004 – – –
Standard Hass 1824.25–1983.525 695.16–2557.375 1381.58–4963.825 7.5–191.615 – – –
Bacon 2160.47 ± 224.41b 418.44 ± 32.19e 337.83 ± 67.42d 0.265 ± 0.22a – 1.74 ± 0.78bc 1.28 ± 0.61bc
Bernecker-43 431.84 ± 48.45e 146.89 ± 48.22e 32.09 ± 21.45d 17.67 ± 9.73a – – 4.95 ± 1.69bc
Dade-3 2092.52 ± 90.78bc 128.39 ± 40.22e 43.43 ± 22.04d 21.12 ± 6.23a – 3.11 ± 058bc 2.30 ± 0.64bc
Day 7354.20 ± 540.88a 5248.19 ± 553.09a 2577.58 ± 401.44b 11.63 ± 5.13a 0.54 ± 0.35ab 0.81 ± 0.42c 11.37 ± 2.25b
FL Hass 1183.90 ± 165.48de 3749.56 ± 722.67b 2905.43 ± 319.65b – 0.25 ± 0.25b 8.80 ± 2.43a 3.50 ± 1.05bc
Lula 1275.08 ± 150.98 cd 2119.21 ± 492.30c 6537.96 ± 798.38a 47.16 ± 47.17a – – 7.80 ± 1.71bc
Miguel 1339.08 ± 236.74 cd 832.54 ± 251.24de 509.42 ± 225.48d 15.08 ± 7.40a – 5.48 ± 1.06ab 1.94 ± 0.63bc
Monroe 1584.03 ± 382.96bcd – – 9.285 ± 6.10a – 0.61 ± 0.43c 3.38 ± 3.15bc
PA-6206 1592.66 ± 254.11bcd 680.63 ± 277.90de 803.39 ± 159.49 cd 20.20 ± 7.70a – 2.91 ± 0.59bc 12.00 ± 2.65b
Pflume 1766.93 ± 231.59bcd 1497.93 ± 455.24 cd 1521.38 ± 306.61c 16.35 ± 4.74a – – –
Simmonds 1881.44 ± 161.96bcd 382.58 ± 61.54e 393.98 ± 132.09d 22.77 ± 8.66a – 0.88 ± 0.60c 0.07 ± 0.07c
Zutano 2312.67 ± 341.60b 66.61 ± 32.55e – – 1.43 ± 0.94a 8.40 ± 3.94a 11.95 ± 5.74b
35706 1172.78 ± 152.73de 9.69 ± 9.70e 6.57 ± 6.57d 14.27 ± 7.87a 0.84 ± 0.52ab 1.15 ± 0.55c 4.81 ± 1.36bc
35707 1103.96 ± 108.25de 177.23 ± 64.46e 12.26 ± 12.27d 16.80 ± 9.15a – – 31.61 ± 9.33a

Within each column, means followed by the same letter are not significantly different at P ≤ 0.05 using the Duncan multiple range test. Values are means ± standard
errors of four replicates of 5 fruits each. Values in Standard Hass were collected from references and were not statistically compared to the experiment data. Data
from 1According to Acree and Arn (2004); Perez-Cacho and Rouseff (2008), or Pereira et al. (2013); 2AT = Aroma threshold; 3Buttery et al. (1988), in water; 4Ahmed
et al. (1978), in water; 5Obenland et al. (2012) and Arpaia et al. (2018).

5
S. Ali, et al.
Fig. 2. Dendrogram clustering of 14 avocado genotypes on the basis of fatty
acids, fatty acid composition ratios and VOCs. The central vertical line shows
Euclidean distance among the clusters and value of Euclidean distance was
about 43%.
fruit of ‘Bacon’ had the lowest concentration of limonene, whereas,
‘Lula’ exhibited the highest value. The sesquiterpenes α-cubebene, α-
copaene and β-caryophyllene, when detected, were present in much
smaller concentrations, except for β-caryophyllene which was greater
than 10 μg kg−1 in ‘Day’ (11.34 μg kg−1), ‘PA-6206’ (12.0 μg kg−1),
‘Zutano’ (11.95 μg kg−1) and ‘35707’ (31.61 μg kg−1) (Table 5).
3.6. Cluster analysis
A cluster analysis showed that avocado genotypes were divided into
three main clusters (clusters were divided on the basis of Euclidean
distance) on the basis of fatty acids, fatty acid groups, fatty acid com-
position ratios, and VOCs (Fig. 2). Cluster I consisted of 11 genotypes:
‘Pflume’, ‘PA-6206’, ‘53707’, ‘Bernecker-43’, ‘Zutano’, ‘Bacon, ‘Monroe’,
‘Miguel’, ‘Simmonds’, ‘Dade-3’, and ‘35706’ (Fig. 2). Cluster II had only
one genotype (‘Lula’), and cluster III contained two avocado genotypes
(‘Day’ and ‘FL Hass’). Among cluster I, ‘Pflume’ had its own sub-cluster
with low TOC, and a high percentage of oleic acid but no sesqui-
terpenes. Another sub-cluster consisted of ‘PA-6206’ and ‘35707’, which
had the highest amount of oleic acid and the lowest amount of palmitic
acid. Likewise, ‘Miguel’, Simmonds’, ‘Dade-3’ and ‘35706’ had the
highest linoleic content. Cluster II singled out ‘Lula’ which had by far
the highest amount of (E)-2-hexenal and limonene, and second to
highest amount of hexanal. ‘Lula’ also had the highest amount of pal-
mitoleic acid, 50–150% more than the other genotypes. Cluster III
contained ‘Day’ and ‘FL Hass’ which had the highest amount of hexanal
and second to highest amount of (E)-2-hexenal following that of ‘Lula’.
‘Day’ also had the highest amount of acetaldehyde. Fatty acids in those
two cultivars were divergent, with palmitoleic acid being high in ‘FL-
Hass’ but low in ‘Day’, and oleic acid high in ‘Day’ and the lowest
(among all genotypes) in ‘FL Hass’.
4. Discussion
Fruit weight, dry matter, and oil content of individual mature
avocado genotypes vary with environmental conditions but show gen-
eral consistency when genotypes are compared in the same environ-
ment. Fruit sizes of Mexican race avocados are reported to be widely
distributed from 75 g to 300 g with high (about 30 %) oil content;
whereas, cultivars of Guatemalan race are reported to have an average
6
Scientia Horticulturae xxx (xxxx) xxxx
weight of 500−600 g with lower oil content (Nakasone and Paull,
1998; Yahia and Woolf, 2011). Dry matter also depends on maturity,
and is usually correlated with oil content: as fruit matures on the tree,
dry matter and TOC increase (Lee et al., 1983; Yahia and Woolf, 2011).
Because oil content does not significantly change after harvest, much
research has been performed since the 1920s to determine harvest
maturity parameters that can be practically used by growers (Lee,
1981). California has established minimum maturity standards based
on fruit dry matter (DM) for a few genotypes including ‘Bacon’ (17.7%
DM) and ‘Hass’ (20.8% DM) (University of California, 2019). Because
there is no such standard established for most genotypes in our study,
we excluded immature fruit that never ripened and only fruit within
acceptable maturity parameters (firmness ≤ 25 N, Pisani et al., 2017),
were analyzed (Table 1). For these genotypes, DM could also be a good
indicator of TOC, as indicated by the high correlation between both
parameters. Furthermore, in spite of the target firmness of 25 N, data
showed a wide range in whole fruit firmness as opposed to mesocarp
firmness (Table 1). It is suggested that whole fruit compression mea-
sures elasticity of the skin, which is quite variable among the genotypes
in this study, while mesocarp firmness (or softness) is an indication of
the eating quality of the fruit and was within acceptable range. Arpaia
et al (2018) defined “eating firmness” at 4.4–6.7 N for ‘Hass’ avocado.
Consistent with classification of avocado races, fruit of ‘Simmonds’
(WI) and ‘Pflume’ (WI) had the lowest oil content, 11.4% and 13.6%,
respectively. However, two other genotypes which are open pollinated
(i.e. only the female parent is known) and in this case, WI seedlings,
‘Bernecker-43’ and ‘Dade-3’, had oil content as high as ‘Hass’, sug-
gesting that the pollen parent were likely not WI. ‘Day’, a G-WI type,
had TOC as low as ‘Pflume’, a WI type. In this study there is not a clear
relationship between the known genetic background and avocado oil
content, likely reflecting our limited knowledge of the pedigrees of
genotypes studied. The highest oil content was for ‘PA-6206’, a G–M
avocado type.
A total of seven fatty acids were detected with varying concentra-
tions. Consistent with previous published reports, the monounsaturated
oleic acid (18:1) was the major fatty acid followed by palmitic (16:0),
linoleic (18:2) and palmitoleic (16:1) acids (Meyer and Terry, 2008;
Ozdemir and Topuz, 2004; Pedreschi et al., 2016; Villa-Rodriguez et al.,
2011; Yanty et al., 2011). Research with ‘Hass’ has shown increasing oil
content as harvest is delayed, with some variability among fatty acid
content, but larger differences in oil content and composition were
reported between growing regions, within a country (Ozdemir and
Topuz, 2004) or across exporting countries (Donetti and Terry, 2014).
For example, percentage of dry weight comprised of oleic and palmitic
acids in ‘Hass’ were respectively: 57–61% and 14–16% for Chilean fruit,
and 40–47% and 23–27% for Peruvian fruit (Donetti and Terry, 2014).
Typically, ‘Hass’ composition is reported with ∼100% and up to
∼300% more oleic acid than palmitic acid (Meyer and Terry, 2008;
Ozdemir and Topuz, 2004; Pacetti et al., 2007). In our study, the gen-
otypes with the highest oleic to palmitic acid ratio (94–97% higher
oleic than palmitic acid) were ‘PA-6206’, a G–M type avocado with the
highest oil content, and ‘35707’, a WI-OP avocado type. On the other
hand, palmitic acid was higher than oleic acid in ‘Bernecker-43’ (WI-
OP) and in ‘FL Hass’ (G–M), and they were in similar levels in ‘Dade-3’
(WI-OP), ‘Simmonds’ (WI), ‘Lula’ (G-WI) and ‘35706’ (WI-OP). The re-
maining genotypes had oleic acid content from 35.67% to 43.29% and
palmitic acid levels from 27.00–30.24 %, with varying known race
associations. In this study there is no trend associating fatty acid
composition by known race, except that the WI avocado types, gen-
erally had a lower oleic to palmitic acid ratio, and that overall, oleic
acid was negatively correlated with palmitic acid. There is not enough
data with this study and others to speculate that avocadoes grown
under subtropical conditions may have a lower oleic to palmitic acid
ratio than avocadoes grown under Mediterranean climates (such as the
Donetti and Terry, 2014, study). It could nevertheless be further ex-
plored. Only one other report shows similar results with ‘Barker’, a WI
S. Ali, et al.
avocado type grown in Brazil, in which palmitic acid content by dry
weight was higher (36.39%) than oleic acid (32.66%) (Galvão et al.,
2014).
Linoleic and palmitoleic acids in ‘Hass’ tend to be more constant
than oleic and palmitic acids across studies, varying between 12–15%
for linoleic acid, and 4–9% for palmitoleic acid (Meyer and Terry, 2008;
Ozdemir and Topuz, 2004; Pacetti et al., 2007; Pedreschi et al., 2016).
Experiments reported here found that these fatty acids had higher va-
lues than those published for ‘Hass’ across all genotypes tested: linoleic
acid ranged from 13.02–20.46 %, and palmitoleic acid from 8.15% to
15.34 %, with one genotype, ‘Lula’ having 23.68% palmitoleic acid
(Table 3). Genotype and environmental conditions or even analytical
methods might explain those discrepancies. Besides ‘Lula’ with high
palmitoleic acid, ‘Simmonds’ had a unique fatty acid profile with about
30% oleic, 30% palmitic and 20% linoleic acid content. Ripeness and
total oil content determine eating quality of avocados (Obenland et al.,
2012), but there is limited research that explores the proportions of
various fatty acids and sensory qualities of ripe avocados. Minor fatty
acids found in this study included arachidic acid (C20:0) (2.61–7.84%),
linolenic acid (C18:3) (0.37-0.45%) and eicosenoic acid (C20:1)
(0.53–1.17%), the latter two were found in only three genotypes. Those
fatty acids were reported in similar concentrations in ‘Hass’ and other
cultivars, except for arachidic acid which was at lower levels in earlier
reports on ‘Hass’ (Galvão et al., 2014; Ozedmir and Topuz, 2004;
Pacetti et al., 2007). Stearic acid (C18:0), reported elsewhere (Galvão
et al., 2014; Ozedmir and Topuz, 2004; Pisani et al., 2017), was not
found in this study.
A diet where fats are primarily polyunsaturated fatty acids is re-
commended by the American Academy of Nutrition and Dietetics
(Vannice and Rasmussen, 2014). Avocado with a relatively high ratio of
(M + P)/S qualifies as a healthy source of fats (Villa-Rodríguez et al.,
2011). A diet where fat composition had higher (M + P)/S and P/S
resulted in greater satiety post ingestion, less caloric intake overall and
a lower lipid deposition in the human body and ultimately, may reduce
the risk of low-density lipoprotein, and cardiovascular diseases (Lawton
et al., 2000). In this work, the values of M/S were greater than 1.0 and
(M + P)/S were higher than 40 in all of the genotypes examined.
The VOCs found in the avocado genotypes in this research were
predominantly characterized by aldehydes, sesquiterpenes, and a
monoterpene. No esters, alcohols or ketones were found, contrary to
other studies (El-Mageed, 2007; Pereira et al., 2013; Galvao et al.,
2016), which may be attributed to sampling method in addition to
different cultivars. Aldehydes were the major compounds, which can be
expected since they are derived from the degradation of lipids, and
avocado fruit is a rich source of lipids. Acetaldehyde is the shortest
chain aldehyde, very volatile, and is not necessarily extracted by sol-
vent methods (Galvao et al., 2016; Pereira et al., 2013). It was present
in large amount in these experiments as well as in Obenland et al.
(2012), where static headspace with SPME sampling was used. Acet-
aldehyde was produced in large amount in all genotypes, except ‘Ber-
necker-43’. Acetaldehyde is known to impart a fresh fruit aroma to
many fruit (Perez-Cacho and Rouseff, 2008). Alcohol dehydrogenase
readily metabolizes it into ethanol that may subsequently be used in
production of certain esters (Rudell et al., 2002). Furthermore, acet-
aldehyde may also be metabolized into acetyl-CoA that can lead to the
biosynthesis of mevalonate and monoterpene production (Obenland
et al., 2012). Hexanal, and (E)-2-hexenal were the next most prevalent
VOCs. Hexanal amounts greatly varied by genotype, from concentra-
tions greater than 2000 μg kg-1 in ‘Day’, ‘FL-Hass’ and ‘Lula’, to less than
100 μg kg-1 in ‘35706’, ‘Zutano’ and ‘35707’ and to none in ‘Monroe’.
Both hexanal, and (E)-2-hexenal have a characteristic “green”, grassy
note with a relatively low aroma threshold, and impart fresh avocado
its typical flavor (Obenland et al., 2012; García-Rojas et al., 2016).
Obenland et al. (2012) reported that acetaldehyde significantly in-
creased and hexanal and (E)-2-hexenal decreased with harvest season in
‘Hass’. Fruit in these experiments were harvested over a period of four
7
Scientia Horticulturae xxx (xxxx) xxxx
months, and being a new study under East-central Florida conditions,
harvesting fully mature fruit that would fully ripen after harvest was of
paramount importance.
Limonene was the only monoterpene detected in these analyses,
whereas β-myrcene, β-pinene or cis-β-ocimene have been reported in
other studies (Galvao et al., 2016; Obenland et al., 2012; Pereira et al.,
2013). Limonene imparts a fresh citrus aroma note, however, based on
limonene’s aroma threshold and the concentrations present limonene
has an insignificant effect on avocado flavor. All genotypes had limo-
nene except ‘FL Hass’, ‘Miguel’ and ‘Zutano’. In previous reports, li-
monene was also not detected in ‘Monroe’ (Pereira et al., 2013), and
only detected as traces in three WI or G-WI cultivars (Galvao et al.,
2016), supporting genotype specificity. The sesquiterpenes, α-cube-
bene, α-copaene, and β-caryophyllene were found in varying con-
centrations. The sesquiterpene VOC α-cubebene is described as having
citrus-like fruity aromas, while α-copaene and β-caryophyllene have
spicy, woody-notes (Pereira et al., 2013). The VOC α-cubebene was
detected only in few genotypes and at low levels; whereas α-copaene
and β-caryophyllene were found in the majority of genotypes and at
higher levels. The predominance of α-cubebene and α-copaene among
sesquiterpenes in this research is consistent with previous reports on
avocado fruit VOCs (Pereira et al., 2013; Galvao et al., 2016). However,
β-caryophyllene was not reported in the earlier studies and its detection
in our work could be ascribed to difference in genotypes, production
region, and analytical method used. The sesquiterpenes such as α-cu-
bebene, α-copaene and β-caryophyllene can impart a distinctive ter-
pene/woody note to avocado (Mahendran, 2016).
5. Conclusion
This research highlights that, in complex avocado horticultural
genotypes, the oil content and fatty acids composition of the fruit me-
socarp are difficult to associate with presumed predominant avocado
race. Some WI types did have the lowest total oil content; however,
others which include WI in their pedigrees (‘Bernecker-43’, ‘Monroe’)
had oil content in the same range as some G–M genotypes. Our work
agrees with many studies published with ‘Hass’ or other G or M avocado
types, that oleic acid was the major fatty acid in avocado mesocarp.
However, this study also revealed that palmitic acid could be as high as
or higher than oleic acid in some genotypes. This may be due to gen-
otype characteristics as much as environmental conditions, and it
would be worth exploring further, as oil content and fatty acids com-
position contribute to sensory and health properties of avocado fruit.
VOCs equally contribute to avocado sensory qualities and flavor; the
large amount of aldehydes in the fruit selected in this study contribute
to "fresh", "fruity" and “green” notes. Limonene likely contributes to
fruity flavor, while the sesquiterpenes, present in some but not all
genotypes likely contribute to spicy and terpene/woody notes. This
research enhances the knowledge of newer cultivars, and potential
breeding parents, that may benefit local growers interested in ex-
panding the cultivation range of the commercial cultivars in Florida or
other regions with similar climatic conditions in the world.
Declaration of Competing Interests
None.
Acknowledgements
The first author wishes to thank financial support from Higher
Education Commission (HEC) of Pakistan under International Research
Support Initiative Program (IRSIP) for PhD scholars. Technical assis-
tance from Blesson Mathews, and Carly Franko is gratefully appre-
ciated.
S. Ali, et al.
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