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David E. Kopsell, William M. Randle, and Mark A. Eiteman

Onion (Allium cepa L.) pungency changes during storage. To better understand these flavor changes, seven onion cultivars representing different storage duration, photoperiodic requirement, and flavor intensity were greenhouse grown and the bulbs stored for 3 or 6 months at 5±3 °C, 0.8 to 1.1 kPa vapor pressure deficit. Bulbs were evaluated using high-pressure liquid chromatography quantification for changes in S-alk(en)yl cysteine sulfoxide (ACSO) flavor precursors and γ-glutamyl peptide (γ-GP) biosynthetic intermediates before storage and monthly thereafter. Before and during storage, cultivars differed in total ACSO, (+) S-methyl-L-cysteine sulfoxide (MCSO), trans-(+)-S-(1-propenyl)-L-cysteine sulfoxide (PRENCSO), (+) propyl-L-cysteine sulfoxide (PCSO), S-2 carboxypropyl glutathione (2-CARB), and γ-L-glutamyl-S-(1-propenyl)-L-cysteine sulfoxide (γGPECSO) concentration. During storage MCSO generally decreased while PRENCSO increased in concentration for most cultivars. The linear increase in PRENCSO concentration during storage was accompanied by a linear decrease in γGPECSO concentration. While not measured in this study, these trends indicate γ-glutamyl transpeptidase activity throughout bulb storage. γ-Glutamyl transpeptidase was previously reported to be active only in the later stages of bulb storage or during bulb sprouting. Changes in ACSO and γ-GP compounds during storage did not follow previously reported changes during storage for enzymatically formed pyruvic acid (EPY) for these cultivars. To better understand what causes flavor changes in onions during storage, future investigations should include analysis of the enzymes involved in flavor development and ACSO hydrolysis products.

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Dean A. Kopsell and William M. Randle

Four cultivars of onion (Allium cepa L. `Primavera', `Granex 33', `Pegasus', and `Sweet Success') were grown to maturity in modified nutrient solutions with or without 2.0 mg·L-1 Na2 SeO4 (1.51 mg·L-1 SeO4 -2). Selenium did not affect total flavor precursor content (ACSO) in `Granex 33', `Pegasus', and `Sweet Success'. However, Se affected several individual ACSOs and precursor intermediates. Selenium decreased γ-L-glutamyl-S-(1-propenyl)-L-cysteine sulfoxide and trans(+)-S-(1-propenyl)-L-cysteine sulfoxide content in all four cultivars. (+)-S-Methyl-L-cysteine sulfoxide content was higher while (+)-S-propyl-L-cysteine sulfoxide content was lower with the added Se for two cultivars. Selenium lowered total bulb S content in all cultivars, and increased the percentage of total S accumulated as SO4 -2 in three cultivars. The effect of Se on the flavor pathway was similar to that found when onions were grown under low S-concentrations. Flavor changes can be expected when onions are grown in a high Se environment, however, specific changes may be cultivar dependent.

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William M. Randle, Jane E. Lancaster, Martin L. Shaw, Kevin H. Sutton, Rob L. Hay, and Mark L. Bussard

Three onion (Allium cepa L.) cultivars were grown to maturity at five S fertility levels and analyzed for S-alk(en)yl-L-cysteine sulfoxide (ACSO) flavor precursors, γ-glutamyl peptide (γ-GP) intermediates, bulb S, pyruvic acid, and soluble solids content. ACSO concentration and composition changed with S fertility, and the response was cultivar dependent. At S treatments that induced S deficiency symptoms during active bulbing, (+)S-methyl-L-cysteine sulfoxide was the dominant flavor precursor, and the flavor pathway was a strong sink for available S. As S fertility increased to luxuriant levels, trans(+)-S-(1-propenyl)-L-cysteine sulfoxide (PRENCSO) became the dominant ACSO. (+)S-propyl-L-cysteine sulfoxide was found in low concentration relative to total ACSO at all S fertility treatments. With low S fertility, S rapidly was metabolized and low γ-GP concentrations were detected. As S fertility increased, γ-GP increased, especially γ-L-glutamyl-S-(1-propenyl)-L-cysteine sulfoxide, the penultimate compound leading to ACSO synthesis. Nearly 95% of the total bulb S could be accounted for in the measured S compounds at low S fertility. However, at the highest S treatment, only 40 % of the total bulb S could be attributed to the ACSO and γ-GP, indicating that other S compounds were significant S reservoirs in onions. Concentrations of enzymatically produced pyruvic acid (EPY) were most closely related to PRENCSO concentrations. Understanding the dynamics of flavor accumulation in onion and other vegetable Alliums will become increasing important as the food and phytomedicinal industries move toward greater product standardization and characterization.

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Marilyn H.Y. Hovius, Irwin L. Goldman, and Kirk L. Parkin

Breeders have found field screening for white rot (Sclerotium cepivorum Berk.) resistance in onion (Allium cepa L.) to be unreliable since consistently moderate to high disease levels that significantly differentiate cultivars do not occur over field sites and years. The objective was to determine if differences in onion white rot resistance levels were associated with differing S-alk(en)yl-l-cysteine sulfoxide (ACSO) levels. A collection of onion breeding lines and hybrids were evaluated in field trials at six sites in 1999-2001. High performance liquid chromatography was used to analyze ACSOs in onion plant organs. Four main cysteine-sulfoxides exist in Allium L. species: methyl (MCSO), 2-propenyl (2-PeCSO), 1-propenyl (1-PeCSO), and propyl (PCSO). 1-PeCSO was predominant in onion leaves, bulbs, and roots. 2-PeCSO was found in trace amounts in onion leaves and roots. There was significantly more 2-PeCSO and total ACSO (roots only) and 1-PeCSO (roots and bulbs) in accessions that were more susceptible to white rot in the field trials. This is the first report of significant differences in ACSO contents among white rot susceptible and resistant onions. A covariance analysis was used to determine if the ACSO levels that significantly distinguished among accessions could predict field onion white rot reaction. 1-PeCSO from both roots and bulbs was the best predictor of field disease incidence in field sites that had low, moderate, and high disease levels. Although the ACSO concentrations were not assessed on an individual plant basis, breeders may be able to screen onions for resistance to S. cepivorum by comparing onion root or bulb 1-PeCSO levels based on the results from this research. White rot incidence in the field should be higher in those plants whose roots and bulbs have the highest levels of 1-PeCSO.

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Pai-Tsang Chang and William M. Randle

Onion is classified as a salt-sensitive crop, though it is found in production on saline soils around the world. While onion flavor intensity has been studied in response to various growing conditions, little is known about its response to salt stress. To understand if NaCl affects growth, flavor development, and mineral content in onion, `Granex 33' plants were grown to maturity with six different concentrations of NaCl ranging from 0 (control) to 125 mm in nutrient solutions. NaCl affected onion fresh weight and altered onion flavor intensity and quality. Plants did not survive the 125 mm NaCl treatments and are not included in the results. As bulb Na+ and Cl- content increased in response to increasing NaCl concentrations, leaf and bulb fresh weight of mature plants decreased. Total bulb S content also decreased with increasing NaCl solution concentrations, while bulb SO4 2- content increasing linearly, indicating that less S was entering the S metabolic stream. Though bulb soluble solids content was not influenced by NaCl concentrations, pungency increased, but only at the highest NaCl concentration. Total flavor precursors and methyl cysteine sulfoxide content increased in response to NaCl, but only at the 100 mm treatment. 1-Propenyl cysteine sulfoxide was generally unresponsive to the salt treatment. Propyl cysteine sulfoxide content decreased then increased in responses to increasing NaCl levels, but was found as a minor flavor precursor. Peptide intermediates measured in the pathway leading to 1-propenyl cysteine sulfoxide and propyl cysteine sulfoxide decreased linearly with increasing NaCl exposure. While NaCl affected onion flavor in this study, severe reductions in growth would prevent onion production under similar saline conditions. For practical purposes, the effects of NaCl on flavor are, therefore, minimal.

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William M. Randle and Rachel Snyder

Mild onion consumption is increasing in the U.S. The ability to produce mild onions depends on selecting proper cultivars and growing them in an appropriate environment. A major decision in producing onions with mild flavor is determining when to stop applying sulfate to the crop. While adequate sulfur is necessary for good early onion growth, high levels of sulfur increase bulb pungency. A study was conducted where sulfate was eliminated from the fertility program at biweekly intervals during onion growth and development. Mature bulbs were then analyzed for flavor precursors and their biosynthetic intermediates, and pungency. Pungency linearly increased from 3.7 to 5.1 μmols pyruvic acid from the earliest cut-off date to the latest cut-off date, respectively. While total milligrams of flavor precursors did not significantly change in response to sulfate elimination, the methyl cysteine sulfoxide: 1-propenyl cysteine sulfoxide ratio did. Methyl cysteine sulfoxide concentration decreased in a quadratic manner while 1-propenyl cysteine sulfoxide linearly increased as sulfate fertility was extended in the growing season. Changes in individual precursors will significantly affect flavor perception as well as flavor intensity.

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Timothy W. Coolong and William M. Randle

The effects of temperature and developmental age on flavor intensity and quality were tested by growing `Granex 33' onions (Allium cepa L.) at 16.5, 22.1, 26.7, and 32.2 (±0.4) °C for 50 days and to maturity. Plants were harvested and evaluated for growth characteristics. Bulbs were then analyzed for sulfur (S) assimilation and flavor development parameters. Total bulb S increased linearly with temperature regardless of bulb age. Bulb sulfate changed little over temperatures, indicating that organically bound S increased with temperature. Total pyruvic acid content (pungency), total S-alkenyl cysteine sulfoxide (ACSO) content and individual ACSOs increased linearly in response to temperature when measured at the two developmental stages. Though trans-(+)-S-(1-propenyl)-L-cysteine sulfoxide was the predominant ACSO at most temperatures, (+)-S-methyl-L-cysteine sulfoxide accumulation was greatest among the individual ACSOs in mature bulbs grown at 32.2 °C. Additionally, (+)-S-propyl cysteine sulfoxide was present in the least amount at all treatment levels and developmental stages. Gamma glutamyl propenyl cysteine sulfoxide and 2-carboxypropyl glutathione peptides in the flavor biosynthetic pathway also increased linearly with temperature. When ACSOs were assessed in onion macerate as a measure of alliinase activity, levels of degraded ACSOs increased linearly with growing temperature. The relative percentage of most ACSOs hydrolyzed, however, did not change in response to growing temperature. This suggested that the activity of alliinase was proportional to the amounts of flavor precursors synthesized. Growing temperature, therefore, should be considered when evaluating and interpreting yearly and regional variability in onion flavor.

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Timothy W. Coolong and William M. Randle

To determine the extent to which sulfur (S) and nitrogen (N) fertility interact to influence the flavor biosynthetic pathway in onion (Allium cepa L.), `Granex 33' onions were grown in hydroponic solution culture with varying levels of S and N availability. Plants were grown at 5, 45, or 125 mg·L-1 sulfate (SO4 2-), and 10, 50, 90, or 130 mg·L-1 N, in a factorial combination. Total bulb S, total and individual flavor precursors and their peptide intermediates in intact onion tissue were measured. To measure the effect of S and N on alliinase activity, flavor precursors were also measured in onion macerates. Sulfur and N availability in the hydroponics solution interacted to influence all flavor compounds except S-methyl-L-cysteine sulfoxide. Levels of S-methyl-L-cysteine sulfoxide were influenced by N and S levels in the solutions; however, no interaction was present. At the lowest SO 4 2- or N levels, most precursors and peptides measured were present in very low concentrations. When SO 4 2- or N availability was adequate, differences among flavor compounds were small. Results indicated that S fertility had a greater influence on trans-S-1-propenyl-L-cysteine sulfoxide (1-PRENCSO) accumulation, while N availability had a greater influence on S-methyl-L-cysteine sulfoxide levels. Flavor precursors remaining in the onion macerates revealed that the percentage of intact precursors hydrolyzed by alliinase were not significantly influenced by either SO 4 2- or N levels in the solutions, except for 1-PRENCSO, which was affected by N levels. Nitrogen and S fertility interacted to influence the flavor biosynthetic pathway and may need to be considered together when manipulating onion flavor compounds.

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W.M. Randle

To test the effects of high nitrogen (N) fertilization levels on onion quality and bulb flavor, `Granex 33' onions (Allium cepa L.) were greenhouse grown in hydroponic solution culture with increasing N concentrations. Nitrogen was adjusted in the solutions with NH4NO3 and increased incrementally from 0.22 g·L-1 to 0.97 g·L-1 over five treatments. Plants were harvested at maturity and subjected to quality, flavor, and mineral analysis. As solution N increased, bulb fresh weight and bulb firmness decreased linearly. Gross flavor intensity, as measured by enzymatically developed pyruvic acid (EPY) increased linearly for N concentrations between 0.22 and 0.78 g·L-1, but EPY was reduced slightly in bulbs grown at the highest N level (0.97 g·L-1). Soluble solids content was unaffected by solution N concentration. Solution N had an affect on flavor quality. Methyl cysteine sulfoxide, which gives rise to cabbage (Brassica L. sp.) and fresh onion flavors upon eating, generally increased in concentration as solution N increased. 1-Propenyl cysteine sulfoxide, which imparts heat, mouth burn, pungency, and raw onion flavors increased between the two lowest N concentrations, and then decreased as solution N increased. Propyl cysteine sulfoxide, which imparts fresh onion and sulfur flavors upon eating, generally increased with increasing solution N concentration. Several minerals were also affected by solution N concentration. Total bulb N and NO3 - increased linearly while B, Ca, and Mg decreased linearly. Total bulb S and K increased and then decreased quadratically in response to increasing solution N. Nitrogen fertility can have a pronounced affect on onion flavor and as a consequence, needs to be considered when growing onions for specific flavor quality and nutritional attributes.

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Artemio Z. Tulio* Jr., Yoshinori Ueda, Hiroyuki Yamanaka, Yoshihiro Imahori, Kazuo Chachin, and Artemio Z. Tulio* Jr.

The emission of methanethiol (MT) and dimethyl disulfide (DMDS) from homogenate fractions of fresh and frozen broccoli tissues was analyzed using gas chromatography coupled with flame photometric detector after incubating for 2 h at 30 °C in a water bath. Both sulfur compounds were detected in the headspace of the residue fraction of fresh broccoli but not frozen tissues. Only DMDS was formed in the filtrate and supernatant fractions of fresh tissues but their emission was also suppressed in frozen tissues. Phosphate buffer treatment reduced the amount of MT formed on the residue of fresh tissues, whereas treatment of enzyme co-factor, pyridoxal phosphate, and its substrate, S-methyl-L-cysteine sulfoxide, in the residue fraction of frozen broccoli did not induce the formation of MT except for DMDS. Both compounds were also inhibited in the residue fraction of the fresh tissues by aminooxyacetic acid, a potential inhibitor of pyridoxal phosphate-dependent enzymes, indicating that these objectionable odors were produced upon the action of cysteine sulfoxide lyase. This enzyme, which yielded strong activity in the residue upon extraction with buffer containing Triton X-100, is highly likely to be a bound enzyme. Inhibition of MT and DMDS in frozen broccoli tissues is likely attributed to the retardation of the enzyme action due in part to the loss of the co-factor and its substrate, and owing to the solubility of MT and its affinity to gaseous condition.