Evolution of Soluble Sugars During Ripening of Papaya Fruit and its Relation to Sweet Taste

M. GOMEZ, F. LAJOLO, AND B. CORDENUNSI ABSTRACT: Fruit ripening is closely associated with compositional and structural changes which can occur before or after harvesting. In papaya fruit, the period of sugar synthesis and accumulation remains poorly understood. The correlation between soluble sugar content and sweetness during papaya ripening was investigated in this study. Soluble sugars accumulated mainly when the papaya fruit was still attached to the plant. After harvest, however, there was still sucrose synthesis, and the sucrose-phosphate synthase activity was highly correlated to the sucrose content, indicating the importance of this enzyme in the process. Sensory analysis showed that there was a dissociation between sugar content and sweet sensory perception, while the pulp softening showed high correlation with the sweetness process, probably due to the easier release of cellular contents in the fully ripened tissue. Keywords: papaya, sensory analysis, soluble sugars, sweet taste

Introduction
fructose contents, which are often used as an index of ripening. Some climacteric fruits, such as green bananas (Cordenunsi and Lajolo 1995) and kiwi fruit (McRae and others 1992), have a high starch content which is metabolized to sucrose after harvesting, leading to the fruit sweetness. In bananas, starch metabolizing enzymes, mainly -amylase and starch phosphorylase along with enzymes related to sucrose synthesis, contribute to this process (Garcia and Lajolo 1988). The accumulation of sucrose seems to be related to sucrose-phosphate synthase (SPS) and sucrose synthase (SuSy) activities. The former increases during banana ripening, while the latter decreases sharply after harvesting (Cordenunsi and Lajolo 1995; Nascimento and others 1997). Other climacteric fruit, such as papayas, do not accumulate starch during development (Paull 1996; Gomez and others 1999). After harvesting and during fruit ripening, sugar changes and sweetness development in papayas are not yet well established, although sweet taste is a possible quality index. Papayas for commercial trade are harvested when the peel color approaches yellow, about 110 d post-anthesis (dpa). At this stage, the pulp is still hard and its taste and flavor (similar to those of carrot) are unacceptable for consumption. Changes in texture, peel, and pulps color, organic acid levels, and synthesis of volatile compounds normally occur during detached papaya ripening, concomitantly with the climacteric period (Paull 1996). There are many contradictions concerning soluble sugar synthesis and accumulation during ripening. Results from investigations done with papayas that ripened attached to the tree, receiving a constant supply of sucrose that originates from photosynthesis in the leaf, are quite different from those obtained in detached fruit. At the initial stages of papaya development, glucose is prevalent among the soluble sugars (Paull 1996). With the changes of coloration in pulp and seeds, starting from the 110th dpa, there is a modification in the sugar profile at which time sucrose becomes the predominant sugar (Chan and others 1979; Selvaraj and others 1982). Chan and others

WEET TASTE IS AN IMPORTANT QUALITY PARAMETER FOR FRUITS. It is usually associated with sucrose, glucose, and

(1979) reported sucrose levels varying from 1.8% on the 110th dpa to 8.0% on the 135 th dpa in attached fruits. On the other hand, a variation in detached fruits of about 2% in the total level of sugars between the 2nd and 22nd days after harvest (dah) was reported by Hubbard and others (1991). Because papayas have a low starch content at harvest time (about 0.1%), it would not be a sufficient carbon source for the increase in sucrose content and for post-harvest sweetening (Gomez and others 1999). Thus, studies are still necessary to improve the understanding of the sweetness process in papayas ripened after harvest. In this study, the changes of soluble sugar contents and in the activities of SPS and SuSy of ripening papayas, after harvesting, were determined. Relationships between these changes, the changes in the sweet taste, and the texture and cell wall composition were also tentatively established.

Materials and Methods

KIN COLOR IS USED COMMERCIALLY TO DESCRIBE PAPAYA MA-

turity (Paull and Chen 1997; Lazan and others 1989; Chan and others 1981). Based on this visual maturity determination, green mature papayas (Carica papaya L. cv. Solo) with about yellowed skin were obtained from a commercial market in So Paulo, Brazil, about 72 h after harvesting. Ethylene and carbon dioxide were measured daily, to ensure that the fruits were in the preclimacteric period. Eight fruit with yellowed skin and hard pulp were placed in a temperature-controlled chamber at 25 C and 85% room humidity (RH), then allowed to ripen naturally.

Sampling related to the ripening period

Papaya harvesting is based on visual determination of skin coloration, resulting in large variation of samples during the maturation stages. In order to avoid these problems, a system to always analyze the same fruit (n = 8) throughout the ripening process was established. A core of fruit pulp was removed daily from the equatorial portion, horizontal to the axis, with the help of a cork borer, 1.8 cm in diameter. Cores of 4 fruit were composite as 1 sub-sample, resulting in 2 subsamples of 4 fruit each. The holes were filled with paraffin

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Ripening of papaya fruits relation to sweet taste . . .

wax and the fruit was returned to the chamber. To avoid contamination, the incision had been previously and locally sanitized with a solution of 5% sodium hypochlorite. The cork borer was sterilized by flaming. The cores were immediately peeled and frozen in liquid nitrogen, and stored in a freezer (-80 C). The sampling procedure was deemed appropriate, because the fruits continued to ripen naturally, displaying the same characteristics that a whole fruit would. The respiration pattern was comparable to that of whole fruit, demonstrating that there was no significant stress due to the sampling method.
Table 1Descriptive definitions of attributes and references used by sensory panel for the evaluation of papaya aroma and flavor at 3 different ripening stages Descriptors Overall aroma Ripe papaya aroma Definition The overall intensity of all papaya odorants as perceived by the nose The intensity of characteristic ripe papaya aroma, perceived by the nose, from a reference of 8 cm3 ripe papaya cubes, with yellow peel and orange and soft pulp The intensity of characteristic green papaya aroma, perceived by the nose, from a reference of 8 cm3 green papaya cubes, with dark green peel and white and hard pulp The perceived intensity of taste of a 10% sucrose solution The intensity of characteristic ripe papaya flavor, perceived by mouth, from a reference of 8 cm3 ripe papaya cubes, with yellow peel and orange and soft pulp The intensity of characteristic green papaya flavor, perceived by mouth, from a reference of 8 cm3 green papaya cubes, with dark green peel and white and hard pulp The perceived intensity of taste of a 2% caffeine solution The perceived intensity of taste of a 5% citric acid solution Force required during mastication of a reference sample of a carrot cube with 1.5 cm3 area

Green papaya aroma

Sampling related to texture and sensory analysis

Texture and sensory analyses were done at only 3 ripening stages (mature green, intermediate, and ripe), using 3 fruit in each stage. At the mature green stage, the fruit had yellowed skin, with hard pulp, as they are normally picked, and were comparable to the ripening control sample at the 3rd day after harvest (dah), as described above. At the intermediary stage (5 th dah), papayas were found with yellowed skin, yellow/orange pulp, and soft but still firm. Ripe papayas (8th dah) showed all the skin yellowed, with totally soft orange/red pulp.

Sweetness Ripe papaya flavor

Green papaya flavor

Ethylene and respiration measurement

For the ethylene and respiration analyses, the fruit was individually placed in a 1 L flask. After 1 h, 1 mL of the air inside the flask was sampled and the composition of gases determined with a gas chromatograph equipped with a thermal conductivity detector for CO 2, and with a flame ionization detector for ethylene (HP 6890 series, Analytical Development Co., HP Instruments, Little Falls, Del., U.S.A.), using a Porapack Q column (Supelco, Bellefonte, Penn., U.S.A.).

Bitterness Sourness Hardness

Carbohydrate analysis
Starch content was determined enzymatically, as described by Cordenunsi and Lajolo (1995). Soluble sugars were extracted 3 times with 80% ethanol at 80 C. The supernatants were combined and the ethanol was evaporated under vacuum. The soluble sugar was properly diluted in pure water and analyzed by high performance liquid chromatogaphy with pulse amperometric detection (HPLC-PAD) (Dionex, Sunnyvale, Calif., U.S.A.), using a PA1 column (Dionex) in an isocratic run of 18 mM NaOH for 25 min. Total soluble sugars were given as the sum of glucose, fructose, and sucrose values. The carbohydrates in the cell wall were extracted with 10% (w/v) water at 90 C, precipitated with 96% ethanol, hydrolyzed with 2 N H2SO 4/121 C for 1 h, then separated by HPLC-PAD on a PA1 column, using the same conditions described above.

cose, 5 mM fructose-6-phosphate, 15 mM glucose-6-phosphate, 15 mM MgCl2, and 1 mM NaF. The freshly desalted enzymatic extracts (60 L) were incubated with 40 L of the medium for 20 min at 37 C. The reaction was stopped with 200 L of 1M NaOH, and the extract was boiled for 10 min to destroy the remaining fructose. The released sucrose was assayed through the thiobarbituric acid method (Percheron 1962). A single unit (U) of both SuSy and SPS activity was defined as the activity producing 1 mol of sucrose or sucrose6-phosphate in 1h/mg protein, respectively.

Enzyme extraction and assay

For the assay of sucrose synthase (SuSy) and sucrosephosphate synthase (SPS) activities, about 1 g of frozen samples were ground in 1 mL of Tris-HCl buffer, 500 mM and pH 7.5, containing 40 mM EDTA, 50 mM ascorbic acid, 15% (v/v) glycerol, and 1 M NaCl. After centrifugation (20 min/20000 g), the supernatant was desalted by a Sephadex G-50 HiTrap (Pharmacia, Uppsala, Sweden), and the enzymatic activities were determined in the protein fraction. SuSy activity was measured in the sucrose synthesis direction with Tris-HCl buffer, 100 mM and pH 7.6, containing 1 mM NaF, 10 mM fructose, 5 mM UDP-glucose, 15 mM MgCl 2. The procedure for the SPS assay in saturated condition was performed with 50 mM Hepes-KOH at pH 7.5, containing 10 mM UDP-glu-

Sensory analysis
Quantitative Descriptive Analysis (QDA), described by Stone and others (1974), was done with samples of green (3 rd dah), intermediate (5 th dah), and ripened (8 th dah) papayas. The sensory analysis was done using a panel of 14 judges selected from among students and employees from the University of So Paulo. These judges were often papaya consumers, but did not have previous experience in sensory analysis of foods. The sessions were carried out under the supervision of a leader, and the analyzed parameters were aroma, flavor, and texture. Initially, the judges developed the descriptive terms of the samples, using the Grid Repertory Methodology (Moskowitz 1983). Under the leaders supervision, there was a discussion of the descriptive terms, which

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Ripening of papaya fruits relation to sweet taste . . .

Table 2Cell-wall neutral sugar content of water-soluble cell-wall polysaccharides of papaya fruit during ripening Days after harvest 3 6 10 Rhamnose 158 183 304 Arabinose 132 136 148 Xylose 56 47 58 Galactose 573 600 389 Glucose 81 33 100

Results are expressed in mg sugar/g of the polysaccharide water-soluble fraction

removed synonyms and unusual terms, and the terms which clearly described similarity or differences between samples were selected. A brief description of each term was developed in agreement and can be seen in Table 1. A File of Sample Evaluation was also developed, associating an unstructured scale of 9 cm to each descriptive term. In order to

define none or strong perception of the analyzed parameters in this scale, the minimum was identified on the left side (0 cm) by none and the maximum was identified on the right side (9 cm) by strong. The judges were instructed to localize the parameter intensity of the sample in the scale, and this position was read as the distance in centimeters from the left anchor. Samples were provided to the judges in plastic dishes with a 3-digit code number. Training sessions were conducted and the sample references that were provided for the judges training can also be seen in Table 1. Finally, each judge made the evaluation in 3 replications, using the developed Files of Sample Evaluation. With the results generated by each judge, an individual ANOVA (variation sources: samples, replication) of each descriptor with the F levels of significance for each sample was calculated (p 0.05), as well as for replication (p 0.05). Judges with consensual averages with the sensory team in more than 80% of the judged attributes were chosen to compose the descriptive sensory panel. The selected team evaluated the 3 samples in 3 replications and the results were analyzed by ANOVA (sources of variation sample, judges, and the sample x judges interaction) and Tukeys average, p 0.05. For statistical analysis, the Statistica 5.0 Stat Soft (Tulsa, Okla.,U.S.A.) was used.

Texture evaluation
Texture analysis was done using a texturometer TA-XT2, equipped with a TA-7 USDA Warner-Bratzler knife with triangle cut (SMS Godalming, Surrey, England) to determine the force of shearing disks of 4-cm diameter, removed from the equatorial region of the fruit. For each maturation stage, 3 fruit were analyzed, and 4 replications for each fruit were done.

Results and Discussion

Profiles of soluble carbohydrates during papaya ripening
Papaya fruit utilized were harvested at a preclimateric point, as indicated by the ethylene and CO2 evolution during ripening (Figure 1A). Starch content was about 0.13% in green fruit (3 dah) and about 0.06% in ripened fruit (8 dah),

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Figure 1Respiration, ethylene production, total soluble sugar (TSS), sucrose (suc), and hexose variation during ripening of papaya fruit. Respiration (Figure 1A, open squares) showed a peak at 6th dah; however, the ethylene production peak (Figure 1A, closed squares) had occurred at the 5th dah. Total soluble sugars (Figure 1B, open triangles) decreased on the 6th dah, due to a decrease in sucrose levels (Figure 1C, closed triangles), while glucose (Figure 1D, closed circles), and fructose (Figure 1D, open circles) remained practically unchanged during the ripening period.

Figure 2A typical HPLC-PAD chromatogram of ripening papaya fruit. It can be seen that no other sugar besides glucose, fructose, or sucrose is present in the sample. In this way, the reported soluble sugars are the only sweetness contributors.

Ripening of papaya fruits relation to sweet taste . . .

Table 3Average values* of flavor, aroma, and texture attributes of papaya fruit, at 3 different stages Aroma Overall aroma Green Intermediary Ripe 1.76B 6.58A 7.69A Ripe papya 0.68B 6.35A 7.77A Green papya 6.31A 1.08B 0.98B Sweetness 1.65B 6.50A 7.48A Flavor Ripe papya 0.70B 6.80A 7.72A Green papya 7.35A 0.56B 1.22B Bitterness 1.81A 0.66A,B 0.50B Sourness 0.69A 0.44B 0.58B Texture Hardness 8.19A 1.84B 1.18B

n =14 judges. Values with different superscripts indicate significant differences (p 0.05) in columns. *The results express the distance, in centimeters, from the left anchor of the scale.

Table 4Contents of soluble sugars (mg/g fresh weight) in 3 ripening stages of papaya fruit Total soluble Sucrose sugar (13.9 5.0)B (26.6 8.1)A (29.8 4.0)A 38.6A 43.8A 48.6A

Glucose 2.8)A

Fructose

Mature green (17.8 (6.9 1.2) A Intermediary (11.8 2.54)B (5.4 1.1) A Ripe (12.7 2.33)B (6.1 1.1) A

Values with different superscripts indicate significant differences (p 0.05) in columns. Results are expressed as means of 5 replicates standard deviation.

which is in agreement with data obtained by Selvaraj and others (1982), and Gomez and others (1999). These data confirmed that papaya fruit does not have enough starch to be hydrolyzed for sweetness production during post-harvest ripening, which suggests that there is an alternative carbon source for sugar synthesis. Total soluble sugars (Figure 1B) varied from 9.5% in green stage (3rd dah) to 10% in the 4th dah, decreasing to 9% in ripe stage (8th dah). These results were very similar to those de-

scribed by Chan and others (1979) for the Solo cultivar, and higher than the contents found by Hubbard and others (1991), which was about 6% for a non-specified cultivar. Sucrose levels (Figure 1C) decreased from about 5% during the ripening period to 2% at the 6th dah, concomitantly with the respiration peak. The same results were obtained in 3 different experiments, suggesting that there is sucrose de novo synthesis during ripening of detached papaya. Glucose and fructose contents (Figure 1D) were very similar to each other and varied from 2 to 3%. The HPLC profiles showed that only fructose, glucose, and sucrose are found in all stages of papaya ripening (Figure 2). The carbon for the sucrose synthesis in papayas may come from the cell wall, which contains about 30% cellulose, 30% hemicelullose, 35% pectin, and 5% proteins (Brett and Waldron 1996). The largest change in the cell walls composition, related to the ripening of many fruits, is the loss of significant amounts of neutral sugars, especially galactose and arabinose (Pressey 1983). The water-soluble polysaccharides from papaya cell wall were analyzed at 3 different ripening stages. Rhamnose content increased, while the arabinose and xylose contents remained practically constant, as shown in Table 2. The main change observed was a decrease in ga-

Figure 3Sucrose synthase (SuSy), sucrose-phosphate synthase (SPS) activities, and sucrose content during papaya ripening. SuSy activity (A, closed squares) showed little correlation to sucrose content (A, open triangles), while SPS activity (B, closed triangles) showed relation to sucrose content (B, open triangles) during the ripening period.

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Ripening of papaya fruits relation to sweet taste . . .

lactose and an increase in glucose contents (increase of the glucose/galactose ratio). These data indicate a degalactosylation of the main polysaccharide chain, an event also observed in apples, strawberries, tomatoes, and in germinating seeds (Brett and Waldron 1996, Pressey 1983). This galactose must be quickly metabolized, since we have not found free galactose in ripe papaya (data not shown). According to these results, galactose could be a source of carbon for sucrose synthesis and would support increased sucrose consumption during the climacteric respiration. Sucrose synthase activity increased about twice during papaya ripening (Figure 3A). It started with 3.5 U in green papayas (3 dah) and finished with 6 U in overripe papaya (10 dah). This behavior was quite different from that obtained for SuSy from banana, whose activity decreased along ripening (Cordenunsi and Lajolo 1995). The reason for this increase is unknown, since no correlation between SuSy activity and sucrose content was found. This enzyme seems to be important for sucrose translocation and utilization when the fruit is still on the tree (Cordenunsi and Lajolo 1995). On the other hand, the profile obtained for SPS activity during papaya ripening followed the tendency of the sucrose content profile (Figure 3B), since the ratio SPS activity/sucrose is constant. The average activities were very similar to those obtained by Hubbard and others (1991), who showed a value around 5 U, which might correspond to a ripe fruit, but they did not show a profile of enzymatic activity. These results support the view that there is sucrose synthesis during detached papaya ripening which may be related to SPS activity. mediate, and ripe stages for aroma, flavor, acidity, and texture. In general, the judges were not able to differentiate the papayas of the intermediate stage from those of the ripe stage, for any parameter. However, there were statistically significant differences among these 2 stages and the mature green stages in all parameters. The mature green samples were clearly different from the other 2 stages concerning sweetness. When the main soluble sugars (glucose, fructose, and sucrose) responsible for the sweet taste were quantified in the same samples used in the sensory analysis (Table 4), no differences were observed between stages. There were no significant differences between total soluble sugar of green (38.6 mg/g FW ), intermediate (43.8 mg/g FW ), and ripe (48.6 mg/g FW ) stages. No other soluble sugars were detected in those samples by means of HPLC-PAD chromatograms, as can be seen in Figure 2.

Sensory analysis and texture

Firmness of samples from the 3 stages were determined by means of a texturometer (Figure 4), and the results obtained were similar to those reported by the sensory panel: there were no differences between intermediate and ripe fruit, and the green fruit showed statistically higher values. According to these results, the sweeter taste of ripe and intermediate fruit could be associated with a softer texture, which would facilitate mastication and liberation of the sugars for the sensory perception. In order to verify this hypothesis, the same papaya samples analyzed by sensory analysis in the 3 stages were cut into small pieces and homogenized in an Ultra-Turrax for different periods (30, 60, 90, and 120 s) to simulate mastication. After being filtered, the homogenate was analyzed by

Sensory analysis and the sugar levels in different stages of papaya ripening
The sensory panel of 14 selected judges developed 9 descriptive terms to describe the sensory attributes of aroma, flavor, and texture of the papayas in the 3 characteristic stages of sweetening. Table 1 shows the descriptive terms generated in agreement, as well as their definitions. Table 3 presents the results for papayas in the mature green, inter-

Sensory and Nutritive Qualities of Food

Figure 4Texture and sweetness of 3 ripening stages of papaya fruit. Both instrumental (open circles) and sensory (open triangles) measurements are in agreement, showing a decrease in the intermediary and ripe stages. Sweetness (closed squares), which showed a negative correlation with hardness, had an increase in the intermediary and ripe fruits

Figure 5Hexose release depending on homogenization time and ripening stage of papaya fruit. Mature green fruit (closed squares) showed strict relation between homogenization time and hexose release. Intermediary (closed circles) and ripe (closed triangles) fruit showed maximum hexose release even at the shorter time (30 s).

Ripening of papaya fruits relation to sweet taste . . .

HPLC-PAD (Figure 5), and showed that, in fact, the sugar liberation did suffer pulp firmness interference. Sucrose was not found among sugars liberated by this procedure, since no care was taken in the process to avoid invertase activity (Gomez and others 1999; Chan and Kwok 1975). For green samples, the liberation of hexoses was highly dependent on the homogenization time. However, even for extended periods (120 s), only one-third of the amounts of hexose liberated from ripe papayas was detected. On the other hand, homogenization time was not important for the intermediate and ripe papayas, since the hexose liberation was practically total at the shortest homogenization time (30 s). It can be concluded that the sweet taste may be related to the loss of papaya pulp firmness, since there was a possible correlation between mastication time and the sweet taste perception in papaya pulp. Flavor is also dependent on the aromatic substances formed in the fruit. Perception of aroma was also analyzed. Papayas in the mature green stage were described as having a characteristic green papaya aroma, clearly different from that of the papayas in intermediate and ripe stages, which were described as having a pronounced aroma of ripe papaya. It should be considered that during the sweetness process, there are changes in the profiles and levels of phenolic compounds, which contribute for the flavor (Paull 1996), and also the disappearance of the latex, which had been considered by the panelists as bitter and a characteristic of green papaya (Table 3). Papaya in the mature green stage were noted to be more acidic than those in intermediate and ripe stages by sensory analysis. These results are in disagreement with those obtained by Arriola and others (1980), Camara and others (1993), and DInnocenzo and Lajolo (2001), who found a slight increase in the titratable acidity with maturation. The low contribution to the flavor offered by acidity (Paull 1996), and this effect, can be explained by the suppression of the acid perception by the rising perception of sweetness in the intermediate and ripe samples. tion from the papaya cells in the mouth during mastication.

References
Arriola MC, Calzada JF, Menchu JF, Rolz C, Garcia R. 1980. Papaya. In: Nagy S, Shaw PE, editors. Tropical and subtropical fruits: composition, properties, and uses. Westport, Conn.: AVI. P 316-340. Brett C, Waldron K. 1996. Physiology and biochemistry of plant cell walls. London:Unwin Hyman. 255 p. Camara MM, Diez C, Torija ME. 1993. Changes during ripening of papaya fruit in different storage systems. Food Chem 46:81-84. Chan Jr HT, Hibbard K, Goo T, Akamine EK. 1979. Sugar composition of papayas during fruit development. Hort Sci 14(2):140-141. Chan Jr HT, Kwok SCM. 1975. Importance of enzyme inactivation prior to extraction of sugars from papaya. J Food Sci 40:70-771. Chan Jr HT, Tam SYT, Seo ST. 1981. Papaya polygalacturonase and its role in thermally injured ripening fruit. J Food Sci 46:190-191. Cordenunsi BR, Lajolo FM. 1995. Starch breakdown during banana ripening: sucrose synthase and sucrose-phosphate synthase. J Agric Food Chem 43:347351. DInnocenzo M, Lajolo FM. 2001. Effect of gamma irradiation on softening changes and enzymes activities during ripening of papaya fruit. J Food Biochem 25:425-438 Garcia E, Lajolo FM. 1988. Starch transformation during banana ripening: the amylase and glucosidase behavior. J Food Sci 53(4):1181-1186. Gomez MLPA, Lajolo FM, Cordenunsi BR. 1999. Metabolismo de carboidratos durante o amadurecimento do mamo (Carica papaya L. cv. Solo): influncia da radiao gama. Cinc Tecnol Aliment 19(2):246-252. Hubbard NL, Pharr DM, Huber SC. 1991. Sucrose-phosphate synthase and other sucrose metabolizing enzymes in fruits of various species. Physiol Plant 82:191196. Lazan H, Ali ZM, Liang KS, Yee KL. 1989. Polygalacturonase activity and variation in ripening of papaya fruit with tissue depth and heat treatment. Physiol Plant 77:93-98. McRae E, Quick WP, Benker C, Stitt M. 1992. Carbohydrate metabolism during postharvest ripening in kiwifruit. Planta 188:314-323. Moskowitz HR. 1983. Product testing and sensory evaluation of foods: Marketing and R & D approaches. Westport: Food and Nutrition Press. 605 p. Nascimento JRO, Cordenunsi BR, Lajolo FM, Alcocer MJC. 1997. Banana sucrosephosphate synthase gene expression during fruit ripening. Planta 203:283288. Paull RE. 1996. Pineapple and papaya. In: Seymour GB, Taylor JE, Tucker GA, editors. Biochemistry of fruit ripening. London: Chapman & Hall. P 302-315. Paull RE, Chen W. 1997. Minimal processing of papaya (Carica papaya L.) and the physiology of halved fruit. Postharv Biol Technol 12:93-99. Percheron F. 1962. Colorimetric determination of sucrose and fructofuranosidase by the thiobarbituric acid reaction. CR Seances Acad Sci 255:2521-2522. Pressey R. 1983. -galactosidases in ripening tomatoes. Plant Physiol 71:132135. Selvaraj Y, Subramanyam MD, Iyer CPA. 1982. Changes in the chemical composition of four cultivars of papaya (Carica papaya L.) during growth and development. J Hort Sci 57:135-143. Stone HS, Sidel JL, Oliver S, Woosley A, Singleton RC. 1974. Sensory evaluation by quantitative descriptive analysis. Food Technol 28(11):24-34. MS 20001589 Submitted 12/7/00, Accepted 7/27/01, Received 7/27/01
This work was supported by Fundao de Amparo Pesquisa do Estado de So Paulo (FAPESP). The authors thank the Coordenadoria Aperfeioamento Pessoal Ensino Superior (CAPES) for research scholarships, and Dr. Maria Aparecida Azevedo Pereira da Silva (UNICAMP) for supervising sensory analysis and for her suggestions during this papers writing.

Conclusions
paya harvest could be derived from galactose, whose levels in the cell wall decreased during fruit ripening. Not sucrose synthase, but sucrose-phosphate synthase activity was highly correlated to sucrose content and seemed to participate in the continuous synthesis of sucrose. The fact that ripe and intermediate papayas are classified as sweeter than green ones, despite the same total soluble sugar content, could be associated to changes in texture, which would result in different sugar libera-

HE CARBON SOURCE FOR THE SUCROSE SYNTHESIS AFTER PA-

Authors Gomez, Lajolo, and Cordenunsi are with the Departamento de Alimentos e Nutricao Experimental, Faculdade de Ciencias Farmaceuticas, Universidade de Sao Paulo, Av. Lineu Prestes, 580-CEP 05508-900-SP Sao Paulo-Brazil. Direct inquiries to author Cordenunsi (E-mail: Hojak@usp.br).

Sensory and Nutritive Qualities of Food