Abbreviations: ASG, acylated steryl glycoside; DGDG, digalactosyldiacylglycerol; FS, free sterols; GL, galactolipids; GlyL, glycolipid; MGDG, monoga-lactosyldiacylglycerol; NL, neutral lipid; PA, phosphatidic acid; PC, phosphatidylcholine; PE
Álvaro Fernández-Cuesta, Ossama Kodad, Rafel Socias i Company, and Leonardo Velasco
. Vegetable oils and oil-based products are the richest dietary sources of phytosterols followed by cereal grains, cereal-based products, and nuts ( Piironen et al., 2000 ). Sterols in foods exist as free sterols, fatty acid esters, steryl glycosides, and
Bruce D. Whitaker
Altered metabolism of membrane lipids has been proposed as a mechanism for the beneficial effects of postharvest calcium treatment on apple quality. A previous study showed that after transfer of apples stored 6 months at 0C to 20C, calcium-treated fruit exhibited slower loss of galactolipid and altered levels of sterol conjugates. The present study of lipids in “control” fruit was conducted as a prelude to further in-depth analyses of the effects of postharvest calcium and heat treatments on lipid metabolism in apples during and after cold storage. Neutral lipid, glycolipid (GL), and phospholipid (PL) fractions were obtained by column chromatography followed by TLC separation of GL and PL classes. The major GL were steryl glycosides (SG), acylated steryl glycosides (ASG), cerebrosides (CB), and mono- and digalactosyl diacylglycerols. Phosphatidylcholine (PC) > P-ethanolamine (PE) > P-irositol (PI) were the major PL. The fatty acids of PC and PE were quite similar, whereas those of PI were more saturated. CB included only 2-hydroxy fatty acids. Among the steryl lipids, free sterols > SG > ASG, with beta-sitosterol >90% of the total sterol in each.
Bruce D. Whitaker
A previous study of lipids from pericarp tissue of tomato fruit ranging from mature-green to red-ripe showed a large increase in total sterols accompanied by dramatic changes in sterol composition and conjugation with ripening. This study was conducted to determine whether similar changes occur in microsomal membranes derived from tomato fruit pericarp. Acylated steryl glycoside (ASG), the predominant steryl lipid, declined during ripening, with increases in steryl glycoside (SG) and free sterol (FS). Only minor changes in fatty acid composition were associated with the drop in ASG. The stigmasterol:sitosterol ratio increased throughout ripening, but much more in Fs than in SG or ASG. The ratio of FS to phospholipid (PL) increased with ripening. However, FS was never greater than 10 percent of the total membrane sterol (TMS), and TMS:PL actually declined over the middle stages of ripening. It is not known why tomato tissues maintain such high levels of ASG and SG, but sterol conjugation is thought to regulate the physical properties of cell membranes.
Bruce D. Whitaker
Plastids and microsomal membranes were isolated from pericarp tissue of mature green and red-ripe tell pepper fruit harvested from greenhouse and field grown plants. The lipid composition of these membrane fractions changed far more with ripening of field grown than greenhouse grown fruit. Also, the phospholipid (PL), free sterol (FS), steryl glycoside (SG) and acylated steryl glycoside (ASG) content of microsomes and plastids from both green and red fruit were very different under the two growing conditions. Total steryl lipids (TSL = FS + SG + ASG), and the TSL/PL ratio, increased in microsomes and decreased in plastids with ripening. These changes were much greater in field grown fruit. The ASG/SG ratio decreased with ripening in both membrane fractions, under both growing conditions. Ripening and growth conditions affected the phospholipid and sterol composition in plastids much more than in microsomes. Lipid changes associated with the chloroplast – chromoplast transformation were similar in field and greenhouse grown fruit, including an increase in the galactolipid/PL ratio. Future studies will assess how differences in membrane lipid composition affect postharvest storage life of the fruit.
Dana F. Faubion and Adel A. Kader
California-grown `Hass' avocado fruit were stored at 5C, in air or a controlled atmosphere (CA) of 2% oxygen and 5% carbon dioxide. Fruit were evaluated at 0, 2, 4, 6, 8, 10, and 12 weeks, both immediately upon removal from storage and after ripening at 20C. Severe chilling injury (flesh browning) developed in the airstored fruit after 6 weeks, while only moderate symptoms were observed in CA-stored avocado fruit after 12 weeks. Lipid peroxidation breakdown products increased during storage and ripening in both air and CA treatments. Sterols, steryl esters, steryl glycosides, glycolipids, and phospholipids were analyzed. Quantity of acylated steryl glycoside in ripe fruit changed from 34 nmoles initially, to 51 or 27 nmoles after 6 weeks at 5C in air or CA, respectively. Glycolipid fatty acid unsaturation in air-stored fruit decreased with the development of chilling injury. Fatty acid unsaturation in phospholipids (phosphatidylinositol, phosphatidylcholine, phosphatidylglycerol, and phosphatidylethanolamine) of air-stored avocados decreased with the development of chilling injury. CA storage delayed the development of chilling injury and the loss of fatty acid unsaturation.
Bruce D. Whitaker
Lipid composition and pigment content were determined in pericarp of `Pik Red' tomatoes (Lycopersicon esculentum Mill.) that were harvested when mature-green (MG) then ripened for 1 or 14 days at 20C, chilled for 11 or 21 days at 2C, or chilled for 21 days and transferred to 20C for 4 days (rewarmed). During ripening, chlorophyll fell below a detectable level, carotenes increased 100-fold, phospholipids (PLs) dropped ≈20%, and galactolipids (GLs) dropped ≈35%. Fatty-acid unsaturation decreased slightly. Steryl esters (SEs), more than free sterols (FSs) and steryl glycosides (SGs), increased at the expense of acylated steryl glycosides (ASGs), and in all four steryl lipids, the stigmasterol: sitosterol ratio rose dramatically, whereas the level of isofucosterol fell sharply. During chilling, chlorophyll declined ≈40% and carotenes ≈60%. PL content did not change, whereas GL fell ≈15%. Fatty-acid unsaturation increased slightly. FS, much more than SG and SE, increased at the expense of ASG. The stigmasterol: sitosterol ratio changed little in ASG, SG, and SE but declined in FS. Isofucosterol increased in FS and SE. Rewarming had little effect on the levels of chlorophyll, carotenes, or PL levels, but caused GL to fall another ≈15%. Fatty-acid unsaturation decreased slightly in GL and ASG. The distribution of total sterol in ASG, SG, FS, and SE changed dramatically, yielding proportions close to those in unchilled MG fruit. Also, 4 days after rewarming, the stigmasterol: sitosterol ratio had increased sharply, particularly in FS and SE, and there was a further rise in isofucosterol in all four steryl lipids. These results indicate that chloroplast damage occurs during chilling, but PL-rich cell membranes are not degraded, even upon rewarming. Changes in sterol composition and conjugation during chilling and after rewarming could result in membrane dysfunction.
R.E. McDonald, T.G. McCollum, and E.A. Baldwin
Mature-green `Sunbeam' tomatoes (Lycopersicon esculentum Mill.) were treated in varying order with C2H4, 42 °C water for 1 hour, 38 °C air for 2days, held 2 days at 20 °C (partial ripening), or not treated and then stored at 2 °C (chilled) for 14 days before ripening at 20 °C. Heat-treated fruit stored at 2 °C and transferred to 20 °C ripened normally, while 63% of nonheated fruit decayed before reaching the red-ripe stage. Partially ripened fruit developed more chilling injury, were firmer, were lighter, and were less red in color than fruit not partially ripened. Lycopene content and internal quality characteristics of fruit were similar at the red-ripe stage irrespective of sequence of C2H4 exposure, heat treatment, or a partial ripening period. Of the 15 flavor volatiles analyzed, 10 were reduced by storage at 2 °C, Exposure to C2H4 before the air heat treatment reduced the levels of four volatiles, while C2H4 application either before or after the water heat treatment had no effect on flavor volatiles. Two volatiles were decreased and two were increased by partial vipening, Storage at 2 °C decreased the level of cholesterol and increased levels of campesterol and isofucosterol in the free sterol pool. Exposure to C2H4 before or following heat treatments, the method of heat treatment, and partial ripening had little effect on free sterols, steryl esters, steryl glycosides, or acylated steryl glycosides in the pericarp of red-ripe fruit. A shortor long-term heat treatment of mature-green tomatoes could permit storage at low temperatures with little loss in their ability to ripen normally, whereas partial ripening did not reduce chilling injury.
Bruce D. Whitaker, Joshua D. Klein, and William S. Conway
Postharvest heat treatment of apples maintains fruit firmness and reduces decay during storage. Four days at 38C are beneficial, but 1 or 2 days are detrimental. The cellular basis of these effects may involve changes in cell wall and membrane lipid metabolism. Lipids from hypodermal tissue of `Golden Delicious' apples were analyzed after 0, 1, 2, or 4 days at 38C. Major lipids included phospholipids (PL), free sterols (FS), steryl glycosides (SG), and cerebrosides (CB). Galactolipids (GL) were minor components. PL content fell ?10% after 1 day at 38C, was unchanged after 2 days, and began to rise again after 4 days. PL class composition did not change with heating, but fatty-acid unsaturation declined throughout. FS and CB content and composition changed little, whereas SG content cropped by ≈20% over 4 days. GL fell ≈50% during 1 day at 38C, with no change at days 2 or 4. A burst of PL catabolism followed by recovery of synthesis may in part explain the different effects of 1-, 2-, or 4-day heat treatments. GL loss (in plastids) may be related to the effect of heat on fruit color (yellowing).
R.E. McDonald, W.R. Miller, and T.G. McCollum
Irradiation is being evaluated as a quarantine treatment of grapefruit (Citrus paradisi Macf. `Marsh'), but it can cause damage to the fruit. Research was conducted to determine if preirradiation heat treatments would improve fruit tolerance to irradiation as they improve tolerance to low temperature injury and to determine if canopy position influenced fruit tolerance to irradiation. Initially, grapefruit were irradiated at 0 or 2.0 kGy at a dose rate of 0.14 kGy·min-1 and selected biochemical changes were monitored over time. There was a marked increase in phenylalanine ammonia-lyase (PAL) activity following irradiation. Maximum activity (≈18-fold increase) was attained 24 hours after irradiation. Subsequently, grapefruit were harvested from interior and exterior canopy positions and irradiated at 0 or 1.0 kGy at a dose rate of 0.15 kGy·min-1 before storage for 4 weeks at 10 °C. Following storage, pitting of flavedo was the most evident condition defect noted as a result of irradiation. Pitting was observed on 27% and 15% of irradiated exterior and interior canopy fruit, respectively, whereas there was no pitting on nonirradiated fruit. Heat treatment before irradiation decreased susceptibility of fruit to damage. Pitting was 26%, 19%, and 17% when fruit were held 2 hours at 20 (ambient), 38 or 42 °C, respectively. Irradiation-induced PAL activity was reduced by temperature conditioning at 38 or 42 °C. Exterior canopy fruit flavedo contained higher levels of total phenols, including flavanols and coumarins compared with interior canopy fruit. Deposition of lignin was not related to canopy position. Neither irradiation nor heat treatment had an effect on total phenols or lignin deposition. Generally, cholesterol, campesterol, stigmasterol, β-sitosterol, and isofucosterol were found to be higher in four steryl lipid fractions in exterior canopy fruit compared with interior canopy fruit. Irradiation increased campesterol in the free sterol and steryl glycoside fractions and decreased isofucosterol in the free sterol fraction. Heat treatments had no effect on individual sterol levels. It seems that irradiation causes a stress condition in the fruit, which leads to pitting of peel tissue. Heat treatment before irradiation reduced damaging effects of irradiation.