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Using Density Measurements to Study the Effect of Excision, Storage, Abscisic Acid, and Ethylene on Pithiness in Celery Petioles

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Using Density Measurements to Study the Effect of Excision, Storage, Abscisic Acid, and Ethylene on Pithiness in Celery Petioles J. AMER. SOC. HORT. SCI. 121(1):137–141. 1996. Using Density Measurements to Study the Effect of Excision, Storage, Abscisic Acid, and Ethylene on Pithiness in Celery Petioles Mikal E. Saltveit and Mary E. Mangrich Mann Laboratory, Department of Vegetable Crops, University of California, Davis, CA 95616-8631 Additional key words, aerenchyma, lightly processed Abstract. The density of excised 2-cm celery (Apium graveolens L.) petiole segments was highly correlated with a subjective evaluation of pithiness. Loss of density and the appearance of pithiness was stimulated by lengthening the duration of storage, raising the storage temperatures above 0C, and excising petiole segments. Segments excised from the upper two- thirds of the petiole lost less density during storage than segments excised from the bottom third of the petiole. Segments with initial high densities lost slightly less density during storage at 5C for 5 weeks than segments that were initially less dense. The extent of pithiness development varied significantly among six cultivars held at 5C for 2 weeks. Treating whole petioles with 1 µM abscisic acid for 4 days significantly increased density loss. Exposing petiole segments to up to 100 µl·liter-1 ethylene in humidified air for up to 2 weeks at 5C did not significantly change density over air controls. The loss of density and the development of pithiness in lightly processed celery petioles could be reduced by selecting resistant cultivars, monitoring water stress during growth, using only segments excised from the upper two-thirds of the petiole, and selecting segments with initial high densities. Air spaces that form in the cortex of many plant tissues such as and Pressman (1980) also found that flooding and nutrient defi- celery petioles, corn roots, tomato stems, and roots of wetland ciency lead to pithiness, but not as rapidly as water deficit. species are collectively called aerenchyma. Aerenchyma develops Ethylene production, stimulated by water deficit, has also been through the lysis of parenchymatous cells of the cortex and is thought of as a mechanism leading to pithiness (Pressman et al., stimulated by a variety of stresses, such as deficiencies in water, 1984). In corn roots, a buildup of endogenous levels of ethylene in oxygen, and minerals, and is thought to enhance oxygen transport nonaerated conditions led to aerenchyma formation (Drew et al., within the plant since it is associated with the tolerance of some 1979, 1981; Konings 1982). Aerating the roots with as little at 0.1 crops to anaerobic soils (Drew et al., 1979). When aerenchyma µl·liter-1 ethylene in air promoted aerenchyma formation (Drew et forms in tomato stems and celery petioles, it is often referred to as al., 198 1). The effect of ethylene was enhanced in nitrogen- or pithiness. Pithiness is characterized by the appearance of whitish phosphate-starved roots (He et al., 1992). The presence of silver regions and air spaces within the tissue, and is a major source of ions, previously shown to inhibit ethylene action in a number of quality loss and decreased shelf life in celery. tissues (Cookson and Osborne, 1978; Saltveit et al., 1978), sup- Several climatic and storage factors have been associated with pressed aerenchyma formation in ethylene-treated and nonaerated the development of pithiness in field-grown celery plants, in roots (Drew et al., 1981). harvested stalks and processed petioles. Abscisic acid (ABA) may Mechanical perturbation induced pithiness in celery and tomato have a role in aerenchyma development since an increase in stems (Pressman et al., 1983, 1984). Lightly processing harvested endogenous levels of ABA followed water stress and the develop- celery also appears to exacerbate pithiness development and limits ment of aerenchyma (Livne and Vaadia, 1972). Water stress the use and shelf life of cut petiole segments. It has been suggested during growth favors development of pithiness in celery (Aloni that these stresses may induce pithiness through their effect on and Pressman, 1979). They found that the level of free ABA in ABA and ethylene. Applying ethephon, a chemical that breaks water stressed celery increased before the onset of pithiness and down to release ethylene, to celery plants in the field stimulated was related to the development of pithiness in celery petioles. parenchyma breakdown and induced pithiness, with the effect Withholding water increased the level of ABA 7-fold after 24 h, more pronounced in the basal region (Pressman et al., 1984). with a rapid increase in pithiness occurring after 3 days. Also, Cellulase appears to be involved in aerenchyma development in exogenously applied ABA induced pithiness in detached petioles, sunflower stems (Kawase, 1979) and maize roots (He et al., 1994). with the degree of pithiness increasing with ABA concentration. The objectives of the research reported in this paper were to They hypothesized that ABA production induced by water stress increase our understanding of the biological processes involved in stimulated the development of pithiness. Aloni and Pressman quantifying, predicting, and controlling the development of pithi- (198 1) subsequently investigated the relationship between water ness in lightly processed celery. We developed an objective stress in the field and ABA content on the development of pithiness technique to measure pithiness based on density measurements of in tomato stems. The degree of pithiness was greater in drought- excised petiole segments and used this technique to examine the stressed plants treated with 20 µM ABA than in water-stressed effects of ABA and ethylene on the development of pithiness in plants alone. Nonstressed plants treated with ABA exhibited more harvested and lightly processed celery petioles. pithiness than controls, but less than water-stressed plants. Aloni Materials and Methods Received for publication 30 May 1995, Accepted for publication 2 I Sept. 1995, The cost of publishing this paper was defrayed in part by the payment of page Plant material and storage conditions. Horticulturally mature charges. Under postal regulations, this paper therefore must be hereby marked celery stalks were obtained from local wholesalers and stored at advertisement solely to indicate this fact. 2.5C until used. Individual petioles were removed from the stalk J. AMER. SOC. HORT. SCI. 121(1): 137–141. 1996. 137 and either used as entire petioles or as excised 3- or 10-cm-long segments. Only stalks and petioles free of external defects were used. After excision, the stalks or segments were placed in 16-liter glass jars that were ventilated with a flow of humidified, ethylene- free air at 0, 5, 10, 15, or 20C. After 1 and 2 weeks, stalks were taken out of the jars, the petioles removed from the stalks, and 2-cm sections excised from the center of the top, middle, and bottom regions for density determination. In one study, some petioles were cut into 10-cm segments and stored for an additional week, after which density measurements were made of the middle 2 cm of randomly selected segments. In another study, three 3-cm seg- ments were held in 25 × 100-mm-diameter plastic petri dishes for 2 weeks. All experiments were repeated with similar results. Subjective determination of pithiness. A freshly made trans- verse cut was used to examine the petiole for visual signs of pithiness. Initially, many petioles were cut and arranged in order of increasing pithiness based on a subjective visual evaluation of each Fig. 1. Relationship between subjective measurement of pithiness and the density segment. The following scale was derived from these initial of petiole segments, Pithiness was rated from 0 (no pithiness) to 9 (severe observations of the development of whitish areas in the petiole and pithiness). a score of 0 to 9 was given to petioles based on the extent of the whitish discoloration. A score of 0 = no discoloration; 3 = a few white areas in several locations; 6 = white areas near most of the vascular strands and/or small air spaces; and 9 = large white areas and/or many air spaces. Density determination. Celery petiole segments were trimmed to the central 2 cm before being weighed to the nearest milligram. Segments were then pierced with a thin 3-cm-long stainless-steel pin. A stand was constructed that held the end of the pin and could be used to lower and completely immerse the segment in a beaker of water on a balance. The increase in weight upon immersing the petiole segment in the water equals the weight of the displaced water. The volume of the celery segments was calculated from this weight measurement by dividing the weight of the displaced water by the density of water. The temperature of the water was measured and the density of water obtained from a table of critical values. The density of the petiole segment was calculated by dividing the weight of the petiole by the calculated volume. ABA applications. Petioles were cut from stalks and placed in 600-ml beakers containing 100 ml of 0, 1, 10, and 100µM ABA. Fig. 2. Effect of temperature and storage duration on the density of excised 3-cm The beaker and tissue were placed at 20C under constant dim petiole segments. The vertical bar represents the LSD value (P = 0.05). fluorescent light. The solutions were replaced as needed. After 96 h, 2-cm segments were excised from the middle of the top, middle, and bottom third of the petioles. The density of the petiole segments was determined as described above. Ethylene exposure. Excised 3-cm celery petiole segments were placed in 25 × 100-mm-diameter plastic petri dishes in 16-liter glass jars containing concentrations of 0.0, 0.1, 1.0, 10, 100, and 200 µl·liter-1 ethylene in air. An ethylene scrubber was included in the ethylene-free jar.
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