Changes in PH, Acids, Sugars and Other Quality Parameters During Extended Vine Holding of Ripe Processing Tomatoes
Changes in PH, Acids, Sugars and Other Quality Parameters During Extended Vine Holding of Ripe Processing Tomatoes
Received: 2 December 2010 Revised: 5 January 2011 Accepted: 6 January 2011 Published online in Wiley Online Library:
Abstract
BACKGROUND: Two important quality attributes of processing tomatoes are pH and titratable acidity. These and other quality
attributes can be affected by tomato fruit maturity and over-maturity. We have determined the magnitude of these maturity
effects in four processing tomato cultivars commonly grown in California.
RESULTS: Allowing tomatoes to remain on the vine for up to 4 weeks after ripening resulted in an increase in fruit pH of between
0.01 and 0.02 unit per day for the four cultivars examined. The increase in pH was paralleled by a decrease in titratable acidity,
due to a loss of citric acid. Glucose and fructose concentrations also declined with increasing maturity after ripening. Other
quality parameters (color, lycopene, total pectin, pectin solubility, and Bostwick consistency) all showed little change.
CONCLUSION: Vine holding of ripe fruit adversely affects quality, especially pH and titratable acidity. Recent problems with
high tomato juice pH encountered by tomato processors in California could be the result of increased average fruit maturity at
harvest.
c 2011 Society of Chemical Industry
In all tomato cultivars examined, pH increased as the fruit ripened       were hand harvested weekly beginning 1 week after tagging for
from the green to pink to red stage, and continued to increase as         a total of 5 weeks. Only undamaged fruit with minimal sunburn
the red ripe fruit remained on the vine. TA was at its maximum            were included in the harvest.
at the beginning of the ripening process then decreased as the               The second trial was grown at the University of California
fruit reached the ripe stage and continued to decrease with               West Side Research and Extension Center in Fresno County,
over-maturity. Soluble solids have been shown to increase during          California. The same four varieties as in the first trial were grown
ripening then remain constant with over-maturity.7 Contradictory          from transplants and watered with subsurface drip irrigation
results have been obtained for the effect of maturity on juice            and standard commercial practices. The first harvest was at
consistency where it has been shown that Bostwick values either           125 days after transplanting with three additional harvests at
increase5,6 or decrease with maturity.9                                   1-week intervals. For each harvest four 10-foot sections of the
   Citric acid is the most abundant acid in tomatoes and the              rows were selected at random and the entire mass of tomato fruit
largest contributor to the total TA.10,11 The decrease in TA with         in that section collected. This fruit was then sorted by hand into
maturity and over-maturity is generally assumed to be due to a            green, marketable red, and unmarketable damaged fruit (culls)
loss of citric acid, although direct measurements of changes in           and the total mass in each category determined. Yields per acre
citric acid concentrations with maturity have not been reported.          were calculated from these sample sections. A portion of the
Two other acids that contribute significantly to the TA are malic         marketable red fruit was set aside and transported to UC Davis for
and glutamic acid. Malic acid is typically present at only one            juice preparation and analysis.
tenth the level of citric acid, although the ratio of malic to citric
can vary considerably between different tomato cultivars.10,11            Preparation and evaluation of juices
Changes in malic acid levels with maturity and over-maturity              Microwave hot-break juice was prepared from 1300 g samples of
have not been reported. Glutamic acid levels have been shown              tomatoes from the field trials as described previously.13 For each
to increase 10-fold as the fruit ripens from the green to the red         variety at each harvest date three separate juices were prepared.
stage.12 Further changes in glutamic acid with over-maturity have         Juices were analyzed for lycopene, color, titratable acidity, and pH
not been reported. Glutamic acid is also an important contributor         as described.13
to tomato flavor.                                                            Supernatants were prepared from these juices by centrifuging
   Recently, new cultivars of tomatoes have been introduced which         at 15 000 × g for 10 min. These supernatants were analyzed
have a greater ability to resist decay once fully ripened and can         for individual sugars and acids using enzyme kits (R-Biopharm,
thus be left in the field longer after reaching the red ripe stage.       Marshall, MI, USA). It has been shown that acid and sugar levels
These ‘extended field storage’ (EFS) cultivars are desirable for          determined with these kits on crude tomato juice supernatants
the growers and processors because they allow greater flexibility         give excellent agreement with HPLC analysis.14 Soluble solids
in the timing of harvests once the fruit has reached the ripe             contents of the supernatants were determined with an Atago
stage, which helps in coordinating harvests with processing plant         PR-32 refractometer, and phosphate concentrations by the Ames
capacity. Changes in fruit quality with EFS are likely to be similar to   method.15
those seen in older non-EFS cultivars, although this has not been
studied. In this study we have examined both EFS and non-EFS
cultivars to determine the changes in sugars, acids, and pH that          Statistical analysis
occur when ripe tomatoes are allowed to remain on the vine.               Data are presented as means ± SE. An unpaired t-test with equal
                                                                          variance was used to calculate the two-tailed P value to estimate
                                                                          statistical significance of differences between means.
EXPERIMENTAL
Tomato plantings
Two separate field trials were grown. In the first trial, fruit of
                                                                          RESULTS
                                                                          Fruit color and lycopene
uniform maturity were obtained by tagging fruit in the field at the
pink stage, then this tagged fruit was hand harvested weekly for          To assess the ripeness stage of the fruit the color and lycopene
5 weeks. In the second trial the entire mass of fruit from a randomly     content of hot-break juice prepared from the fruit was determined.
selected 10 foot row section was harvested and sorted weekly for          The a/b ratio, a measure of redness, increased between the first
four weeks. Four cultivars of processing tomatoes, Heinz 2401             harvest at 7 days after tagging, and the second harvest at 14 days,
(H2401), Sun Seeds 6368 (N6368), Heinz 9557 (H9557), and AB2              indicating that the fruit was not fully ripe until the second harvest
were examined. Two of these, H2401 and N6368, are considered              (Fig. 1). The lycopene content of the juice similarly did not reach its
EFS cultivars.                                                            maximum level until the second harvest. The general correlation
   Four cultivars of processing tomatoes, Heinz 2401 (H2401), Sun         between lycopene and a/b ratio has been noted previously.16 Both
Seeds 6368 (N6368), Heinz 9557 (H9557), and AB2 were grown                the a/b values and the lycopene content of the juice from N6368
in two separate field trials. Two of these cultivars (H2401 and           fruit were lower than those of the other three cultivars at all stages
N6368) are considered EFS cultivars. The first trial was grown in         of maturity.
Yolo County, California, at the Vegetable Crops Research Field
Station, University of California Davis. Plants were grown from           pH and titratable acidity
transplants and watered with subsurface drip irrigation using             The pH of hot-break juice increased with increasing maturity in all 4
standard commercial practices. At 96 days after transplanting,            cultivars examined (Fig. 2A). The only obvious differences between
fruit at the pink stage were identified and tagged with small             cultivars were the lower pH values for the H2401 tomatoes at all
adhesive stickers. Completely exposed fruit at the top of the             stages of maturity. Both the EFS cultivars (H2401, N6368) and the
canopy was avoided to reduce the incidence of sunburn in the              non-EFS cultivars (H9557, AB2) showed similar pH increases with
tagged set of fruit. A total of 500 of each variety were tagged. Fruit    maturity. The pH increase between the first harvest at 7 days, and
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                                                      c 2011 Society of Chemical Industry                                  J Sci Food Agric (2011)
Tomato fruit maturity and pH                                                       www.soci.org
          A 2.30                                                                      A              4.70
                       2.20
                                                                                                     4.60
                       2.10
                       2.00                                                                          4.50
         a/b
1.90
                                                                                        pH
                                                                                                     4.40
                                               H2401
                       1.80                                                                                                                              H2401
                                               AB2                                                   4.30
                       1.70                                                                                                                              AB2
                                               H9557
                                                                                                     4.20                                                H9557
                       1.60                    N6368
                                                                                                                                                         N6368
                       1.50
                              0   10            20             30         40                         4.10
                                                                                                            0         10             20            30            40
                                       Days after pink stage
                                                                                                                           Days after pink stage
B 120 B 65
                       100                                                                            60
   Lycopene (µg g–1)
                                                                                                      55
                        80
                                                                                      TA (µeq g–1)
                                                                                                      50
                        60
                                                                                                      45
                                                 H2401
                        40                       AB2                                                  40            H2401
                                                 H9557                                                              AB2
                        20                                                                            35
                                                                                                                    H9557
                                                 N6368
                                                                                                      30            N6368
                         0
                              0   10            20             30         40                          25
                                                                                                            0         10             20             30            40
                                       Days after pink stage
                                                                                                                           Days after pink stage
Figure 1. Color and lycopene content of tomato juices prepared from
tomatoes of different maturities. (A) Color measured as the Hunter Lab a/b                C           5.0
ratio. (B) Lycopene content.
                                                                                                      4.9
                                                                                                      4.8
the final harvest at 35 days, varied between 0.29 and 0.31 unit
                                                                                                      4.7
for the four cultivars examined. This is equal to a pH increase of
approximately 0.01 unit per day.                                                                      4.6
   Total TA of the hot-break tomato juices decreased with
                                                                                        pH
                                                                                                      4.5
increasing tomato maturity (Fig. 2B). This decrease ranged from
                                                                                                      4.4
17 µeq g−1 for the H2401 cultivar to 9 µeq g−1 for the AB2 cultivar.
The decrease in TA mirrored the increase in pH. The H2401                                             4.3
tomatoes, which had a notably lower pH than the other cultivars,                                      4.2       y = –0.7703 ln(x) + 7.3559
had the highest TA. A plot of pH versus TA for all 124 juice samples                                                  R2 = 0.8555
prepared in this study showed a clear relationship between pH                                         4.1
and TA (Fig. 2C).                                                                                     4.0
                                                                                                         20         30          40           50           60          70
                                                                                                                                            –1)
                                                                                                                                 TA (µeq g
Organic acids
The rise in pH and decrease in TA indicates that acid concentrations               Figure 2. (A) Effect of maturity on tomato juice pH. (B) Effect of maturity
                                                                                   on titratable acidity (TA). (C) Relationship between pH and TA.
in the fruit are declining with maturity. The predominant acid in
tomatoes is citric acid. Citric acid levels declined with maturity in
all four tomato cultivars (Fig. 3A). Citric acid levels were 22–30%
lower at the final harvest than at the first harvest, for the four                 in all four cultivars then either declined or remained constant at
cultivars examined. As with the pH and TA measurements, the                        later harvests. Phosphate ions also contribute to the measured
H2401 tomatoes were notably higher in acid content than the                        TA. Free phosphate concentrations in these tomato juices ranged
other three cultivars, but showed a similar decline with maturity.                 from 3 to 4 mmol L−1 but showed no change with maturity or any
Glutamic acid levels were lower than citric acid and did not show                  consistent difference between cultivars (data not shown). These
an obvious decline with maturity in any of the cultivars (Fig. 3B).                phosphate concentrations are in line with what has been reported
The change from the first to the last harvest ranged from +12% to                  previously.11
−16%. Malic acid levels were even lower than glutamic and also                        From the measured tomato juice pH, the expected TA of the
did not decline with maturity (Fig. 3C). From the first harvest (day               juice can be calculated from the measured concentrations of
7) to the third harvest (day 22) malic acid levels actually increased              citric acid, glutamic acid, malic acid, and phosphate, and their
     A                      5.0                                                                                65
                                                                            H2401
                            4.5                                             AB2
                                                                            H9557                              55
     Citric acid (g L–1)
                            4.0
                                                                            N6368
                                                                                                Predicted TA
                            3.5                                                                                45
                            3.0
                                                                                                               35
                            2.5
                            2.0                                                                                                               R2 = 0.9703
                                  0     10             20             30            40                         25
                                              Days after pink stage
     B                      2.5                                                                                15
                                                                                                                 15   25      35         45          55           65
                                                                                                                              Measured TA
                            2.0
    Glutamic acid (g L–1)
                                                                                              the second and fifth harvests were small but significant (P < 0.05)
                            1.0       H2401                                                   and ranged from 10 to 12% for the four cultivars. Fruit of the AB2
                                      AB2                                                     cultivar had the highest sugar levels while H2401 had the lowest.
                            0.5       H9557                                                      Soluble solids content, measured with a refractometer and
                                      N6368                                                   expressed in ◦ Brix, is a commonly used measure for assessing
                            0.0                                                               tomato quality. The soluble sugars glucose and fructose are the
                                  0     10             20           30              40
                                                                                              largest contributor to the total soluble solids. Between the first
                                              Days after pink stage
                                                                                              and last harvests soluble solids declined by between 4% and 7%
    C                       0.8                                                               for the four tomato cultivars, in agreement with the measured
                                                                                              declines in glucose and fructose (Fig. 5B). Overall soluble solids
                            0.7
                                                                                              showed a good correlation with the sum of glucose plus fructose
                            0.6                                                               concentration (Fig. 5C).
    Malic acid (g L–1)
                            0.5
                                                                                                 Other quality attributes changed relatively little over the 4 weeks
                                                                                              of field holding. Consistency, measured as Bostwick values,
                            0.4                                                               increased with later harvest times (Fig. 6A). For all four cultivars the
                            0.3                                       H2401                   average Bostwick values at the fifth harvest were higher than those
                                                                      AB2                     from the first harvest (6–15% higher depending on the cultivar) but
                            0.2
                                                                      H9557                   for no individual cultivar was this difference significant at P < 0.05.
                            0.1                                                               Pectin content and composition are known to be important
                                                                      N6368
                            0.0                                                               contributors to the consistency of tomato juice. Between the first
                                  0     10             20           30              40        and fifth harvests changes in total pectin, measured as galacturonic
                                              Days after pink stage                           acid (GalUA), were small (<5%) and not significant at P < 0.05
Figure 3. Changes in acids with maturity. (A) Citric acid. (B) Glutamic acid.                 (Fig. 6B). Similarly, no significant changes were found in the water
(C) Malic acid.                                                                               solubility or the degree of methyl esterification of pectin between
                                                                                              the first and fifth harvests (data not shown). Although no large
                                                                                              change in consistency or pectin content occurred within a cultivar
known pKa values. When the predicted TA was compared with                                     there were obvious differences between cultivars. The cultivars
the measured TA there was a very good correlation between                                     H2401 and H9557 had substantially higher pectin contents and
these two values for all 124 juice samples prepared in this study                             lower Bostwick values than the AB2 and N6368 cultivars. Overall
(Fig. 4). Predicted TA values were equal to 86% of the measured                               Bostwick consistency showed a significant correlation with pectin
TA values on average, indicating that only 14% of the measured                                content (Fig. 6C), which is in agreement with previous research
TA acidity was due to other acids not measured. These other                                   findings.9
acids would include ascorbic, oxalic, as well as numerous amino
acids including aspartic and γ -aminobutyric. In some varieties of
                                                                                              Effects of delayed harvesting
tomatoes these latter two amino acids have been shown to occur
at levels comparable to that of glutamic acid in ripe fruit.12                                By tagging individual fruit it was possible to successively harvest
                                                                                              fruit of uniform maturity to determine the effect of maturity on
                                                                                              acidity and pH. However, this approach does not address the
Sugar and soluble solids                                                                      question of how large a change in pH will occur if an entire field,
Glucose and fructose levels were highest at the second harvest                                containing fruits of varying maturities, is harvested at successively
when the fruit reached full ripeness, then declined slightly over                             later dates. Nor does it allow for determining the effect of extended
the next 3 weeks of vine holding (Fig. 5A). The decreases between                             vine holding on the yield of intact red fruit. To answer these
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                                                                           c 2011 Society of Chemical Industry                                  J Sci Food Agric (2011)
Tomato fruit maturity and pH                                                                           www.soci.org
   A                                                                                                      A                        20
                                30
   glucose + fructose (g L–1)
18
                                                                                                            Bostwick (cm)
                                                                                                                                   16
                                25
                                                                                                                                   14
                                                          H2401
                                                                                                                                   12
                                20                        AB2
                                                          H9557
                                                                                                                                                                 H2401             AB2
                                                                                                                                   10
                                                          N6368                                                                                                  H9557             N6368
                                15                                                                                                  8
                                      0        10               20           30                 40                                       0         10             20           30           40
                                                       Days after pink stage                                                                             Days after pink stage
                                                                                                          B                       5000
  B
                                5.5
                                                                                                                                  4500
                                5.0
                                                                                                                                  4000
                                4.5                                                                                               3500
                                               H2401
                                4.0                                                                                               3000
                                               AB2
                                3.5
                                               H9557                                                                              2500                      H2401              AB2
                                               N6368                                                                                                        H9557              N6368
                                                                                                                                  2000
                                3.0                                                                                                      0        10             20           30           40
                                      0        10                 20           30               40
                                                                                                                                                        Days after pink stage
                                                       Days after pink stage
                                                                                                           C                       23
  C 6.5
                                                                                                                                   21
                                 6
                                                                                                                                   19
                                                                                                           Bostwick (cm)
       Soluble Solids (°Brix)
5.5 17
                                 5                                                                                                 15
                                                                                                                                   13        R2 = 0.6891
                                4.5
                                                                                                                                   11
                                 4
                                                                            R2 = 0.7301                                             9
                                                                                                                                    1500           2500             3500           4500
                                3.5
                                                                                                                                                       Pectin GalUA (µg g–1)
                                 3
                                  15      20             25            30      35          40          Figure 6. (A) Effect of maturity on Bostwick consistency. (B) Effect of
                                                                                                       maturity on pectin. (C) Correlation between Bostwick consistency and
                                               Glucose + Fructose (g L–1)                              pectin content.
Figure 5. (A) Changes in sugar (glucose + fructose) with maturity.
(B) Changes in soluble solids with maturity. (C) Correlation between soluble
solids and sugars.
                                                                                                       yield by the end of the 4-week period. These results show the
                                                                                                       superior field-holding ability of the EFS cultivars.
                                                                                                          Delayed harvesting resulted in juice with a higher pH in all
questions a second field trial with a different protocol was used.                                     varieties. Average pH ranged from 4.38 to 4.49 at the first harvest,
In this trial the same four cultivars of tomatoes were grown and                                       then increased to a range of 4.70 to 4.89 at the final harvest
entire replicate 10-foot sections of rows were harvested in four                                       21 days later (Fig. 8A). The pH of the final harvest was more than
weekly harvests. The yield of marketable red fruit was determined                                      0.2 unit higher than that in the first trial, which may indicate that
and a sample of this red fruit was used to prepare hot-break juice                                     the average maturity at the final harvest in the second trial was
for analysis.                                                                                          beyond the range of maturities examined in the first trial. As in the
   The yield data showed a clear difference between the cultivars                                      first trial the rise in pH was accompanied by a decline in TA due to
(Fig. 7). The two EFS cultivars, H2401 and N6368, had higher yields                                    a loss of citric acid (Fig. 8B and C). Changes in soluble solids and
in all four harvests and showed no decline in yield until the fourth                                   the concentrations of glucose, fructose, glutamic acid, and malic
harvest. In contrast, the yield of the non-EFS cultivars, AB2 and                                      acid were all similar to what was observed in the first trial (data
H9557, dropped steadily with each harvest reaching near zero                                           not shown).
                                       40                                                              A                     5.00
  Marketable Red Yield (tons acre–1)
                                       35                                                                                    4.90
                                                                                                                                       H2401
                                       30                                                                                    4.80      AB2
                                       25                                                                                              H9557
                                                                                                                             4.70
                                                                                                          pH
                                       20                                                                                              N6368
                                                                                                                             4.60
                                       15       H2401
                                                AB2                                                                          4.50
                                       10
                                                H9557                                                                        4.40
                                        5
                                                N6368
                                                                                                                             4.30
                                        0                                                                                       120   125      130     135     140     145      150
                                        120   125       130     135      140      145      150
                                                                                                                                               Days from transplant
                                                        Days from transplant
DISCUSSION 50
                                                                                                       TA (µeq g–1)
The results presented here, in agreement with several earlier                                                                 45
reports, show that tomato juice pH increases and TA decreases
                                                                                                                              40
with extended vine holding after fruit maturity. Hanna8 found that
                                                                                                                              35       H2401
the average increase in pH was about 0.3 unit over 30 days of
vine holding, which is similar to what we found here. Others have                                                                      AB2
                                                                                                                              30
also reported increases in pH and declines in TA with increased                                                                        H9557
tomato maturity.5,6,10 Citric acid is the most abundant organic acid                                                          25
                                                                                                                                       N6368
in tomatoes and the decrease in TA measured with increased fruit                                                              20
                                                                                                                               120    125      130      135     140      145      150
maturity was due to a loss of citric acid from the fruit. During
ripening in tomatoes, as in other fruits, declines in acid levels are                                                                          Days from transplant
accompanied by increases in sugars. At least a portion of this
change may be due to the metabolic conversion of acids into                                            C
                                                                                                                              4.0
sugars by gluconeogenesis.17 Such a conversion of acids to sugars
did not appear to take place during extended vine holding of ripe
                                                                                                       Citric acid (g L–1)
tomatoes because the decline in citric acid was not accompanied                                                               3.5
by increases in the concentration of glucose and fructose. Rather
the loss of organic acids from the mature fruit appears to be
                                                                                                                              3.0
entirely through respiration.                                                                                                         H2401
   EFS and non-EFS cultivars showed no difference in their loss                                                                       AB2
of citric acid with maturity. Nor was there any other obvious                                                                 2.5     H9557
difference between these types of cultivars other than the ability                                                                    N6368
of the EFS fruit to remain intact longer after ripening, allowing
                                                                                                                              2.0
for a greater yield of marketable fruit in later harvests. Thus the                                                             120   125      130     135     140      145       150
delayed harvesting made possible by EFS tomatoes will lead to pH                                                                               Days from transplant
increases and possible problems for processors.
                                                                                                    Figure 8. Increase in pH and decreases in TA and citric acid with later
   The increase in pH with maturity can be compared with the
                                                                                                    harvest date.
increase in pH observed in recent years at processing plants in
California. Data collected on incoming truck-loads of tomatoes
arriving at these processing plants has shown that over the 8 year                                  differences caused by these cultural practices was a change in the
period from 2001 to 2008 the average pH of fruit has increased by                                   relative maturity of the fruit at harvest.18 The difficulties in defining
about 0.01 unit per year for most common cultivars.4 For the four                                   harvest maturity and its relationship to environmental conditions
cultivars examined here the pH increases with delayed harvest                                       and fruit age have been discussed by others.19
were between 0.01 and 0.02 unit per day. Thus the observed                                             The obvious approach for minimizing problems with high pH
increase in average pH could be explained if the average maturity                                   would be to reduce the field holding of ripe fruit as much as
at harvest has been increasing by only 1 day per year. Over this                                    possible. This may not be practical given that the advantages
same time period two new cultural practices have been widely                                        field holding allows for coordinating harvests with processing
adopted: the use of transplants rather than direct seeding of                                       plant capacity. Since the principal change in the fruit causing the
fields, and drip rather than flood irrigation. While it is possible                                 undesirable rise in pH is a loss of citric acid, breeding for higher
that these new cultural practices are, by themselves, affecting                                     initial citric acid content in the fruit, especially in EFS cultivars
fruit pH, it seems more likely that their effect is through changes                                 where increased fruit maturity is more likely to occur, would be
in fruit maturation. This was the case in previous work where it                                    desirable. The higher citric acid content of the H2401 cultivar shows
was shown that differences in fruit quality between conventional                                    that such an approach is possible. Analysis of pH data collected
versus organic production could be explained if one of the principal                                during commercial production has shown that H2401 typically has
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                                                                                 c 2011 Society of Chemical Industry                                              J Sci Food Agric (2011)
Tomato fruit maturity and pH                                                 www.soci.org
the lowest average pH of any of the major cultivars.4 Alternatively,          8 Hanna GC, Changes in pH and soluble solids of tomatoes during vine
the pH in the final product may be reduced by adding citric acid                   storage of ripe fruit. J Am Soc Hortic Sci 78:459–463 (1961).
to the tomato juice during processing. Addition of citric acid has            9 Liu Y and Luh BS, Effect of harvest maturity on free amino acids,
                                                                                   pectins, ascorbic acid, total nitrogen and minerals in tomato pastes.
been routine for decades in California during production of whole                  J Food Sci 44:425–428 (1979).
peeled and diced tomato products. The practice has been less                 10 Stevens MA, Citrate and malate concentrations in tomato fruits:
common in paste production but may become necessary in the                         Genetic control and maturational effects. J Am Soc Hortic Sci
future.                                                                            97:655–658 (1972).
                                                                             11 Paulson KN and Stevens MA, Relationships among titratable acidity,
                                                                                   pH and buffer composition of tomato fruit. J Food Sci 39:354–357
                                                                                   (1974).
ACKNOWLEDGEMENTS                                                             12 Boggio SB, Palatnik JF, Heldt HW and Valle EM, Changes in amino acid
We would like to thank the California League of Food Processors                    composition and nitrogen metabolizing enzymes in ripening fruits
                                                                                   of Lycopersicon esculentum Mill. Plant Sci 159:125–133 (2000).
and its Tomato Research Committee for supporting this project.               13 Barrett DM, Weakley C, Diaz JV and Watnik M, Qualitative and
                                                                                   nutritional differences in processing tomatoes grown under
                                                                                   commercial organic and conventional production systems. J Food
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