Phytochemistry Letters 52 (2022) 33–39

Contents lists available at ScienceDirect

Phytochemistry Letters
journal homepage: www.elsevier.com/locate/phytol

The impact of post-harvest storage on sweet corn aroma

Jessica P. Yactayo-Chang a, Susan Boehlein b, Robert L. Beiriger c, Marcio F.R. Resende Jr. b,
Robert G. Bruton a, 1, Hans T. Alborn a, Maritza Romero a, William F. Tracy d, Anna K. Block a, *
a
US Department of Agriculture, Agricultural Research Service, Chemistry Research Unit, Center for Medical, Agricultural, and Veterinary Entomology, 1700 SW 23rd
Drive, Gainesville, FL 32608, USA
b
University of Florida, Horticultural Sciences Department, 2250 Hull Road, Gainesville, FL 32611, USA
c
University of Florida, Everglades Research and Education Center, 3200 East Palm Beach Road, Belle Glade, FL 33430, USA
d
Department of Agronomy, University of Wisconsin-Madison, Department of Agronomy, 1575 Linden Dr., Madison, WI 53706, USA

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

Keywords: Fresh market sweet corn (Zea mays L.) is a high sugar, high moisture vegetable that, due to its fast metabolic rate,
Aroma is susceptible to rapid post-harvest decay at ambient temperatures. Immediate cooling and storage at low but not
Flavor freezing temperatures are well established industry practices for maintaining post-harvest sugar content. Less is
Post-harvest storage
known about how post-harvest storage temperature impacts other important sweet corn consumer traits, such as
Sweet corn
Volatiles
aroma. In this study we measure the production and stability of the cooked sweet corn aroma compounds
dimethyl sulfide, dimethyl disulfide, dimethyl trisulfide, 3-methyl furan, 3-methyl butanal and 2-methyl butanal.
We reveal that these aroma compounds are also susceptible to post-harvest decay if sweet corn is stored at
ambient rather than cool temperatures. Furthermore, the quantities and post-harvest decay rates of aroma
volatiles vary among different sweet corn lines, indicating that sweet corn genetics can impact maintenance of
post-harvest flavor.

1. Introduction harvest respiration and maintain sugar content (Becerra-Sanchez

Sweet corn (Zea mays L.) is a type of maize with mutations in the
endosperm starch synthesis pathway that prevent the conversion of
sugar to starch, leading to increased sweetness. These include mutations
in either Shrunken2 (Sh2), the large subunit of AGPase, or Sugary1 (Su1)
an isoamylase type DBE (Zhang et al., 2019). Sweet corn with mutations
in sh2 have only sugar in their kernels while sweet corn with su1 mu­
tations have both sugar and starch and can convert kernel sugar to starch
after harvest. This leads to sh2 sweet corn accumulating more sugar than
su1 sweet corn, the sh2 varieties are therefore often called supersweet
corn.
The loss of starch accumulation, coupled with harvest at the milk
growth stage, leads to a high sugar and moisture content in sweet corn
kernels. The resultant juiciness and sweetness that make it such an
appealing food, also make it highly susceptible to post-harvest decay.
Current fresh market sweet corn production practices involving rapid
cooling close to, but not below, zero degrees Celsius act to limit post-
et al., 2021). The maintenance of kernel sugar content correlates with
prolonged sweetness which is an important trait for consumers (Winter
et al., 1955).
Less is known about how temperature and post-harvest storage im­
pacts other factors important for consumer preference in sweet corn
such as aroma. Several aroma volatiles have been identified in cooked
sweet corn including dimethyl sulfide, which gives the distinctive sweet
corn aroma. Other sweet corn aroma volatiles include hydrogen sulfide,
acetaldehyde, methanethiol, ethanol, ethanethiol, 2-acetyl-1-pyrroline,
2-acetyl-2-thiazoline-hydroxy-2-propanone, 2-hydroxy-3-butanone,
and 2,3-butanediol (Bills et al., 1968; Self et al., 1963; Flora et al.,
1974; Buttery et al., 1994). Sweet corn juice contains 1-heptanol,
2-methyl-2-butenal, (Z)-3-nonen-1-ol, 3-ethyl-2-methyl-1,3-hexadiene,
and 2,4-bis(1,1-dimethylethyl)phenol (Feng et al., 2020).
Sweet corn kernel characteristics including sugars and dimethyl
sulfide content have been shown to have genetic variability in different
sweet corn lines (Azanza et al., 1996a). Potential QTLs (Quantitative

Abbreviations: GC-MS, Gas Chromatograph-Mass Spectrometer; 7143, Seminis EX08767143 (Bicolor, Supersweet (Sh2-type)); Su1, Sugary1; Sh2, Shrunken2; QTL,
Quantitative Trait Loci.
* Corresponding author.
E-mail address: anna.block@usda.gov (A.K. Block).
1
Present address: Kaycha Labs Gainesville, 2444 NE 1st Blvd, Suite 700, Gainesville, FL 32609, USA.

https://doi.org/10.1016/j.phytol.2022.09.001
Received 9 June 2022; Received in revised form 31 August 2022; Accepted 2 September 2022
Available online 13 September 2022
1874-3900/Published by Elsevier Ltd on behalf of Phytochemical Society of Europe.
J.P. Yactayo-Chang et al.
Trait Loci) have been identified for dimethyl sulfide production and
sensory panels have been used to identify potential QTLs for some flavor
attributes including corn aroma and grassy aroma (Azanza et al.,
1996b). However, the mechanistic basis for these QTLs and their roles in
aroma remain enigmatic.
As sweet corn is primarily eaten after cooking, the impact of post-
harvest storage on aroma is important both for constitutive aroma
compounds as well as those produced during the cooking process itself.
In this study we assess the impact of post-harvest storage at different
temperatures on the production of sweet corn aroma volatiles and
investigate the impact of genetics on the stability of both aroma and
sugar content in a selection of sweet corn inbreds and populations. Using
our analytical methods we monitored the production dimethyl sulfide,
dimethyl disulfide, dimethyl trisulfide, 2-methyl butanal, 3-methyl
butanal and 3-methyl furan in cooked sweet corn. We observed that
these volatiles are impacted by post-harvest storage temperature,
revealing that sugar is not the only flavor component in sweetcorn that is
lost during post-harvest storage at ambient temperatures. Furthermore,
we revealed that genetic factors in sweet corn can impact the post-
harvest stability of sugars and the aroma volatiles dimethyl sulfide
and dimethyl disulfide. These findings show the untapped breeding
potential for sweet corn with increased “shelf stable” flavor.
2. Material and methods
2.1. Sweet corn material and treatments
Ears from the commercial corn hybrid Seminis EX08767143
(Bicolor, Supersweet (Sh2-type)), from here on referred to as 7143, were
harvested from commercial fields in Belle Glade Florida. The ears were
rapidly chilled on ice and kept on ice for two days. Husks were removed
from the ears and the ears were placed at either 4ºC or 25 ºC until 6, 13 or
20 d after harvest. Samples for sugar analysis were flash frozen it liquid
nitrogen. For volatile analysis, each sample consisted of a 20 g ear
segment that was placed in a steamer plate with 2 mL water and cooked
in a domestic microwave oven (1000 watts) for one minute. Four sam­
ples were analyzed for each treatment.
Sweet corn inbreds: C81, C90, I97, Wh09009, Wh9137R,
Wh10127R, and Wh13064, along with populations Mexican Dent_sh2
and Pease Crosby Sweet corn were grown at the University of Florida/
IFAS Plant Science Research and Education Unit located in Citra, Flor­
ida. The experiment was established in an alpha-lattice design with two
replications. Genotypes were planted in single-row plots with 12 plants
per row. Inbred lines C81 and C90 are su1 lines developed in Con­
necticut (Tracy, 2000). Inbred line I97 contains the sh2 mutation and
was developed by the sweet corn breeding program at University of
Florida. Mexican Dent and Pease Crosby are two open pollinated pop­
ulations, containing the sh2 and su1 mutations, respectively. The inbred
Wh lines contain the sh2 mutation and were developed by the sweet corn
breeding program at University of Wisconsin-Madison. The plants were
self-pollinated and kernels from all lines were harvested 21 d after
pollination and either flash frozen in liquid nitrogen or stored husked for
7 d at 25 ºC before flash freezing. Frozen kernels were used for sugar
analysis. For volatile analysis each sample consisted of 10 g of frozen
kernels that were placed in a steamer plate with 2 mL water and cooked
in a domestic microwave oven (1000 watts) for two minutes. Two
samples were analyzed for each plant and two plants were used for each
treatment giving a total sample size per treatment of four.
2.2. Solventless volatile collection and analysis
Sweet corn samples were transferred to a 125 mL glass mason jar
immediately after steaming. As described in Alborn et al. (2021), the lid
of the mason jar contained two junctions consisting of ¼ inch Teflon
Swagelock connections. One junction was equipped with a
single-tapered splitless liner with a plug of silanized glass wool (Agilent
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Phytochemistry Letters 52 (2022) 33–39
Technologies, Santa Clara, CA) and filled with activated charcoal, and
was used to supply clean air to the jar. The second junction was used to
collect volatiles by attaching a thermal desorption filter consisting of the
same type of single-tapered splitless liners filled with Tenax TA® (60/80
mesh; Supelco, Bellefonte, PA, USA). Fifty mL of air was pulled out of the
jar and through the filter at a constant rate of 10 mL/min by an in-house
designed volumetric syringe pump. The pump contains of a 60 mL
polypropylene syringe coupled to a stepper motor with 200 step­
s/revolution enabling a smooth and consistent draw of air through the
system.
An Agilent 6890 Gas Chromatograph (GC) coupled to an Agilent
5975 Mass Spectrometer (MS) (Santa Clara, CA), was used for analyses
of the volatile collections. The splitless injector was modified with the
addition of a temperature gradient cold trap (Alborn et al., 2018). In that
way, the injector could be used to desorb the filters and the volatiles
efferently focused on the column, thus eliminating the need for flash
heating of the trap. GC-MS was equipped with a Supelco SPB 122–0132
column (DB-1 30 m x 250 μM x 0.25 μM). The injector temperature was
set to 200 ◦ C and splitless injection beginning at a helium carrier flow of
0 psi (for insertion of 140 filter) and then at 0.01 min increased to a
constant pressure of 10 psi. After 2 min the purge valve was opened, and
the carrier gas flow reduced to 0.07 mL/min and a linear velocity of 35
cm/min kept for the remainder of the analysis. The oven was kept at
30 ◦ C for the initial 3 min, followed by ramping at 10 ◦ C/min ramp to
260 ◦ C and kept at that temperature for the last 5 min of the analysis.
Commercial standards were analyzed using the same system and
temperature program but with inactivated cold trap and no initial
injector pressure shut off, thus giving very similar retention times be­
tween the solventless system and solvent injections. The volatile com­
pounds were validated using commercial standards obtained from
Sigma-Aldrich (St. Louis, MO), and analyzed using following mass-to
charge ratio (m/z) and retention times (RT): dimethyl sulfide (m/z 62,
RT 4.84 min), dimethyl disulfide (m/z 94, RT 7.02 min), dimethyl
trisulfide (m/z 126, RT 10.6 min), 3-methyl-butanal (m/z 58, RT 5.96
min), 2-methyl-butanal (m/z 58, RT 6.0 min), and 3-methyl furan (m/z
82, RT 5.58 min).
2.3. Sugar analysis
Sugars were determined using the sucrose, D-fructose and D-glucose
assay procedure from Megazyme (K-SUFRG) scaled down to 96 well
plate format. Briefly, fifteen kernels per sample were lyophilized and
pulverized and approximately 15 mg resuspended in 1 mL of cold water
(4 ◦ C), vortexed, and incubated at 80 ◦ C for 15 min. Solids were removed
by centrifugation at 13,000xg for 10 min and the supernatant was
diluted so measurements could be performed in the linear range of the
assay (0–5 μg). Enzymes, and substrate were added according to the
Megazyme assay procedure but were scaled down to accommodate a 96
well format. Standard curves for glucose, fructose and sucrose were used
to calculate the sugar content in each sample as described in (Finegan
et al., 2022). Briefly, the amount of sucrose was calculated by sub­
tracting the amount of glucose from the amount of sucrose plus glucose
in the sample.
2.4. Statistical analysis
Data were analyzed using the Statistical Analysis System (SAS)
software. Statistical differences at P ≤ 0.05 between treatments were
calculated by one-way Analysis of Variance (ANOVA) followed by a
Tukey’s multiple comparisons procedure.
3. Results
3.1. Post-harvest cold storage slows sugar metabolism in fresh sweet corn
To assess the impact of post-harvest storage temperature on the
J.P. Yactayo-Chang et al.
metabolism in kernels of fresh corn-on-the-cob (variety 7143), sugar
content of the kernels from ears stored at either 4 ºC or 25 ºC was
determined at various time points after harvest (Fig. 1). Independent of
the test storage conditions, the most abundant simple sugar in these
sweet corn kernels is the disaccharide sucrose. The fresh 7143 kernels
had approximately 400 mg/g dry weight of sucrose and this level was
maintained for 20 d post-harvest examined in ears stored at 4 ºC. In
contrast ears stored at 25 ºC displayed a rapid and continuous significant
decrease in sucrose levels over time reaching 100 mg/g by 20 d post-
harvest (Fig. 1A). Sucrose can be broken down into the mono­
saccharides fructose and glucose. These monosaccharides, that were
present at around 30 mg/g and 50 mg/g respectively in the fresh 7143
kernels, were also not significantly impacted by 20 d of post-harvest
Fig. 1. The impact of storage temperature on sweet corn sugar metabolism.
Field harvested ears of the sweet corn hybrid 7143 were packed on ice and two
days after harvest placed at 25 ºC (RT) or 4 ºC. At the indicated time points the
levels of the major sugars (A) sucrose, (B) glucose, and (C) fructose were
determined in cobs. Bars are mean ± standard error, n = 3; *significant dif­
ference levels between the two treatments; p˂0.05 by unpaired t test.
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Phytochemistry Letters 52 (2022) 33–39
storage at 4 ◦ C (Fig. 1B and C). The amounts were, however, signifi­
cantly lower in 7143 kernels after 13 d of post-harvest storage at 25 ºC
and reached less than 10 mg/g after 20 d of post-harvest storage at 25 ºC,
indicating their conversion to starch and phytoglycogen.
3.2. Post-harvest storage at 25 ºC also leads to loss of fresh sweet corn
aroma volatiles
To investigate if post-harvest storage temperature also leads to
changes in aroma volatile production, we collected and analyzed
headspace volatiles from sections of the ears of fresh corn-on-the-cob
(variety 7143) immediately following cooking. Chemical structures of
these compounds are shown in Fig. 2. This analysis revealed the pres­
ence of the high levels of the sulfur containing compounds dimethyl
sulfide (10–50 ng/g/min), dimethyl disulfide (0.05–0.12 ng/g/min)
and dimethyl trisulfide (0.01–0.02 ng/g/min). Furthermore, the alde­
hydes 3-methyl butanal (isovaleraldehyde) and 2-methyl butanal (2-
methylbutyraldehyde) were produced at around 0.2 ng/g/min, while 3-
methyl furan was produced at approximately 2–3 ng/g/min (Fig. 3).
None of the six volatiles measured were significantly impacted by
post-harvest storage for 20 d at 4 ºC (Fig. 3). In contrast, ears stored at 25
ºC displayed a continuous significant decrease in volatile production
following cooking after 13 d of post-harvest storage.
3.3. Sweet corn aroma volatiles in different inbreds and populations
To assess the impact of different genetic backgrounds on the pro­
duction and maintenance of sweet corn aroma volatiles, kernels were
harvested from seven different sweet corn inbreds and two sweet corn
populations grown together in the same environmental conditions.
Dimethyl sulfide levels showed around a three-fold variation between
the different lines and populations, with around 60 ng/g/min in line
C81 and only 20 ng/g/min in line Wh10127R (Fig. 4A). Inbred C81 also
displayed the highest levels of dimethyl disulfide at around 0.19 ng/g/
min, while four-fold lower concentrations were observed in other lines
such as Wh10127R and the population Mexican_Dent_sh2 (Fig. 4B).
The volatile dimethyl trisulfide was highest in line Wh13064 at
around 0.03 ng/g/min and around six-fold lower at around 0.005 ng/g/
min in population Mexican_Dent_sh2 (Fig. 4C). The levels of 3-methyl
furan were similar in all lines tested (Fig. 4D). The levels of 3-methyl
butanal (Fig. 4E) correlated with those of 2-methyl butanal (Fig. 4F)
in the sweet corn inbreds and populations, with 2-methyl butanal the
more abundant of the two. There is a five-fold difference between the
C81 that has the highest amount of these two compounds and Pease
Crosby Sweet corn that has the lowest amounts. These data show that
different sweet corn lines and populations when grown in the same
conditions can produce different quantities of aroma volatiles.
Fig. 2. Sweet corn aroma volatiles.
J.P. Yactayo-Chang et al.
Fig. 3. The impact of storage temperature on sweet corn volatiles. Field har­
vested ears of the sweet corn hybrid 7143 were packed on ice and two days
after harvest placed at 25 ºC (RT) or 4 ºC. At the indicated time points the
production of the major headspace volatiles (A) dimethyl sulfide, (B) dimethyl
disulfide, (C) dimethyl trisulfide, (D) 3-methyl furan, (E) 3-methyl butanal and
(F) 2-methyl butanal were analyzed in cooked cob samples. Bars are mean
± standard error, n = 4; *significant difference levels between the two treat­
ments; p˂0.05 by unpaired t test.
To assess whether genetic variation could also impact the stability of
these aroma volatiles at ambient temperatures, we analyzed the volatiles
of kernels following storage for 7 d at 25 ºC (Fig. 4). The relatively short
post-harvest storage at 25 ºC had no significant impact on dimethyl
trisulfide, 3-methyl furan, 3-methyl butanal or 2-methyl butanal pro­
duction in any of the lines or populations tested. However, dimethyl
sulfide production was significantly reduced in the population Mex­
ican_Dent_sh2 following this treatment (Fig. 4A), and lines C81 and C90
showed significant reductions in dimethyl disulfide production after
post-harvest storage (Fig. 4B). The other lines and populations were not
significantly altered in their production of these volatiles after 7 d of
post-harvest storage. These data indicate that there are genetic factors
underlying the stability of dimethyl sulfide and dimethyl disulfide pro­
duction during post-harvest storage.
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Phytochemistry Letters 52 (2022) 33–39
Fig. 4. Volatile production in kernels of different sweet corns showing the
impact of seven days of post harvest room temperature storage on the volatiles.
Nine sweet corn inbreds and populations (1. C81, 2. C90, 3. I97, 4. Mex­
ican_Dent_sh2, 5. Pease Crosby Sweet corn, 6. Wh09009, 7. Wh09137R, 8.
Wh10127R, 9. Wh13064) were grown in the same field and kernels were
harvested 21 days after pollination (21 DAP). These flash frozen kernels or
kernels from the same ears maintained for seven days at room temperature (21
DAP+7RT) before flash freezing were cooked and head-space volatiles
analyzed. Graphs show quantities of the major volatiles Dimethyl sulfide (A),
Dimethyl disulfide (B), Dimethyl trisulfide (C), 3-methyl furan (D), 3-methyl
butanal (E) and 2-methyl butanal (F). Bars are mean values ± standard error
on the mean, n = 4; *significant difference levels between the two treatments;
P ≤ 0.05 by unpaired t test.
3.4. Sugar levels and post-harvest stability in different sweet corn inbreds
and populations
To assess the impact of different genetic backgrounds on the pro­
duction and maintenance of soluble sugars in sweet corn sucrose,
glucose and fructose content in kernels of the nine different sweet corns
at 21 d after pollination with or without 7 d post-harvest storage at 25 ºC
was determined (Fig. 5). As anticipated the inbreds C81 and C90 and the
population Pease Crosby Sweet corn, which all carry the su1 allele, had
relatively low levels of sucrose, around 20 mg/g (Fig. 5A, B and E),
J.P. Yactayo-Chang et al.
Fig. 5. The impact of seven-day post-harvest storage on sugars in various sweet
corns. Kernels from nine sweet corn inbreds and populations were harvested 21
days after pollination (21 DAP) and flash frozen or harvested and stored for
seven days at 25 ºC before freezing (21 DAP+7RT). Amounts of sucrose (Suc),
glucose (Glu) and fructose (Fruc) were determined for each treatment in the
lines C81 (A), C90 (B), I97 (C), Mexican_Dent_sh2 (D), Pease Crosby Sweet corn
(E), Wh09009 (F), Wh09137R (G), Wh10127R (H) and Wh13064 (I). Bars are
mean ± standard error of the mean, n = 4; *significant difference levels be­
tween the two treatments; P ≤ 0.05 by unpaired t test.
compared to the lines and populations with the sh2 allele. The exception
to this was Mexican_Dent_sh2 that had low sucrose levels despite having
the sh2 allele (Fig. 5D). The sh2 containing inbreds Wh09009 and
Wh09137R had three-fold higher sucrose levels, around 60 mg/g
(Fig. 5F and G). Glucose and fructose contents were significantly lower
than sucrose in all lines.
The materials also varied in the rate of sugar metabolism during post-
harvest storage. For instance, inbred C90 displayed significant decreases
in sucrose, fructose and glucose after seven days of storage at 25 ºC
(Fig. 5B). Inbred I97 only displayed a significant reduction of glucose
content during this treatment (Fig. 5C). In contrast, inbreds C81 and
Wh10127R showed a significant increase in glucose content after post-
harvest storage (Fig. 5A and H). Materials Mexican_Dent_sh2, Pease
Crosby Sweet corn, Wh09009 and Wh13064 showed no significant
impact on sugar content during seven day post-harvest storage (Fig. 5D,
E,F,I). These data show that although su1 and sh2 are major drivers of
sugar content in sweet corn, they are not the only factors involved.
Furthermore, additional genetic factors exist that impact post-harvest
sugar stability.
4. Discussion
The major volatiles observed in cooked sweet corn in this study were
dimethyl sulfide, dimethyl disulfide, dimethyl trisulfide, 3-methyl
butanal, 2-methyl butanal, and 3-methyl furan. Dimethyl sulfide has
been observed in several studies of sweet corn aroma and Self et al.
(1963) found high levels of dimethyl sulfide, hydrogen sulfide, and
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Phytochemistry Letters 52 (2022) 33–39
acetaldehyde and relatively low levels of methanethiol, ethanethiol,
acetone, and methanol in the corn. Bills and Keenan4 found dimethyl
sulfide as well as high levels of ethanol in canned corn but did not
observe the mercaptans reported by Self et al. (1963). Flora and Wiley
(1974) also found hydrogen sulfide, methanethiol, acetaldehyde,
ethanol, ethanethiol, and dimethyl sulfide, while Buttery et al. (1994)
also found 2-acetyl-1-pyrroline, 2-acetyl-2-thiazoline-hydrox­
y-2-propanone, 2-hydroxy-3-butanone, and 2,3-butanediol. The differ­
ences in cooking, sampling, analytical methods and sweet corn varieties
used, as well as growing location and environmental conditions, may
account for the detection of different compounds in the different studies.
Dimethyl sulfide is produced by thermal treatment of the amino acid
S-methylmethionine in many food products (Scherb et al., 2009). Its low
odor threshold of 0.06 ppm, distinctive smell, and high abundance make
it the primary factor for determining the characteristic aroma of cooked
sweet corn. Dimethyl disulfide and dimethyl trisulfide are produced via
thermal degradation of the amino acid methionine, via a methanethiol
intermediate, and temperature pH and antioxidant status can all impact
the rate of formation of these compounds (Pan et al., 2021). Dimethyl
disulfide and dimethyl trisulfide have odor thresholds of 7 ppm and 10
ppt respectively. Even though these sulfur compounds are less abundant
than dimethyl sulfide they could still contribute to sweet corn aroma.
The aldehydes 3-methyl butanal and 2-methyl butanal are derived
from the enzymatic break-down of leucine and isoleucine respectively
(Gonda et al., 2012). In low concentrations 3-methyl butanal gives a
fruity/cocoa aroma and 2-methyl butanal a coffee/nutty/cocoa aroma.
3-methyl furan on the other hand can be made during cooking via the
Maillard reaction from amino acids and reducing sugars, by the by the
lipid oxidation of polyunsaturated fatty acids, and thermal oxidation of
ascorbic acid or carotenoids (Limacher et al., 2008). Different amounts
of these volatile compounds can lead to different aroma and taste pro­
files in sweet corn.
Our study reveals the importance of post-harvest storage at cool
temperatures (4 ◦ C) not only to maintain the levels of kernel sugars but
also those of the aroma volatiles important for sweet corn flavor. As
active metabolism is required for kernel sugars to be converted into
starch (Carey et al., 1984) it is likely that the slowdown of kernel
metabolism is also responsible for the maintenance of kernel aroma
compounds at cool post-harvest storage temperatures.
Several of the aroma compounds are produced from thermal break­
down of amino acids during cooking. Proteins and amino acids are
catabolized as alternate energy sources in stressed, senescent and car­
bohydrate limited plant tissues (Araújo et al., 2011). The higher meta­
bolic rate of kernels at ambient temperature (25 ◦ C) compared to cool
temperatures (4 ◦ C) likely results in increased protein/amino acid
catabolism during storage. The resultant predicted loss of amino acid
precursors could therefore lead to reduced production of aroma
volatiles.
It was previously shown that QTLs exist in sweetcorn for sensory
aromas and for the abundance of dimethyl sulfide (Azanza et al., 1996b),
though the dimethyl sulfide quantification was done on dried kernels
autoclaved with toluene rather than on cooked fresh kernels. In this
study we confirmed that genetic variation in sweet corn can contribute
to dimethyl sulfide production and can also impact other aroma volatiles
including dimethyl disulfide, dimethyl trisulfide, 2-methyl butanal and
3-methyl butanal.
The amino acid methionine, a precursor for dimethyl sulfide,
dimethyl disulfide and dimethyl trisulfide, is the most abundant amino
acid in sweet corn kernels and can vary significantly between different
varieties (Yang et al., 2021). Genetic differences in methionine pro­
duction capacity may impact the production of these three aroma vol­
atiles. The higher amounts of all three volatiles in inbred line C81
compared to line C90 is a potential indicator of such a difference.
Methionine in developing seeds arises from the S-methylmethionine
cycle (Cohen et al., 2017). In this cycle leaf derived methionine is con­
verted to S-methylmethionine by methionine S-methyltransferases, of
J.P. Yactayo-Chang et al.
which there are at least 4 in corn (Ranocha et al., 2001). S-methyl­
methionine is then transported into the developing kernel and converted
back to methionine by homocysteine methyltransferases. Differences in
the enzymes of S -methylmethionine cycle (Cohen et al., 2017) could
therefore also contribute to the observed variations in dimethyl sulfide
production.
The quantities of aldehydes 3-methyl butanal and 2-methyl butanal,
derived from leucine and isoleucine respectively (Gonda et al., 2012),
display correlated changes in production between different genotypes.
High levels of both compounds are observed in inbreds C81 and
Wh09009, while low levels occur in inbred C91 and population Pease
Crosby. In all cases examined the 2-methyl butanal levels are higher
than those of 3-methyl butanal. These data suggest that variations in
shared enzymes in the biosynthesis pathways of these volatiles may be
contributing to the differences seen between different sweet corn in­
breds and populations. Interestingly no significant variation was seen in
the amounts of 3-methyl furan produced, maybe due to its production
during cooking from various precursors.
The sweet corn inbreds and populations analyzed in this study
showed expected variations in kernel sugar content driven, for the most
part, by whether they were su1 or sh2 mutants. The sugar content did
not correlate with differences in aroma volatiles. For instance, the
inbred C81 (su1) has low sugar levels, while the inbred Wh09009 (sh2)
has high sugar levels. These two inbreds had among the highest levels of
aroma volatiles of all the sweet corn in this study. These data indicate
that there is genetic potential in sweet corn to breed for both improved
flavor and improved sweetness characteristics.
Furthermore, we show that genetic variation can also impact the
post-harvest maintenance of sugar and aroma and thus the shelf life of
fresh market sweet corn. Kinetics of sugar loss in the hybrid line 7143
showed sucrose levels decreasing following 6 d post-harvest storage at
ambient temperature. However, of the various lines and populations
examined inbred C90 was the only other sweet corn to show a significant
decrease in sucrose by 7 d storage at ambient temperature. This shows
that the rate of sugar loss is not linked to the total amount of sugars in
the kernel. The inbred C90 was also one of the two lines examined (the
other was C81) that displayed a significant decrease in dimethyl disul­
fide production after post-harvest storage, hinting at a possible high
post-harvest metabolism in this line. Of the aroma volatiles analyzed in
the different sweet corns only dimethyl sulfide and dimethyl disulfide
displayed significant post-harvest reductions after 7 d and the former
only in the population Mexican_Dent_sh2. On the other hand, we saw no
genetic differences in the maintenance of dimethyl trisulfide, 3-methyl
furan, 2-methyl butanal or 3-methyl butanal. There could be several
explanations as to why these compounds do not show variations in their
post-harvest stability. The most likely is as our assay assesses reduced
stability by sampling at a 7 d time point and some variation would be
observed at longer storage times. Alternate explanations include that
these compounds either have less genetic influence on their stability or
the lines we examined did not have significant differences in the loci
involved.
In conclusion, this study showed that there is the genetic potential in
sweet corn for breeding for both improved flavor and improved shelf
life. Both these traits are of value to the consumer as they lead to taster
fresh market sweet corn.
Funding
This work was funded by the United States Department of Agricul­
ture, USA grant numbers USDA-NIFA-SCRI 2018–51181–28419 and
USDA-ARS 6036–11210–001–000D.
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
38
Phytochemistry Letters 52 (2022) 33–39
the work reported in this paper.
Data availability
Data will be made available on request.
Acknowledgments
The use of trade name, commercial product or corporation in this
publication is for the information and convenience of the reader and
does not imply an official recommendation, endorsement or approval by
the USDA or the Agricultural Research Service for any product or service
to the exclusion of others that may be suitable. USDA is an equal op­
portunity provider and employer. The authors declare no conflicts of
interest with this study.
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