HortScience 21(5): 1165-1167. 1986.

Parthenocarpic cucumbers (iCucumis sati­

vus L. cv. Deliva) of known age were ob­
Effect of Stage of Development on tained from a commercial greenhouse near
Lodi, Calif. In 1982, fruit were harvested

Postharvest Behavior of Cucumber and divided into 10 developmental stages

based on size, weight, and color. In 1983,
freshly opened flowers were tagged to iden­
Fruit tify fruit of known age. Since fruit previ­
ously set exert an inhibitory effect on the
Angelos K. Kanellis1, L.L. Morris, and M.E. Saltveit, Jr. growth of subsequent fruit (14), those fruit
Department of Vegetable Crops, Mann Laboratory, University of present on the vine at the time of tagging
were removed. Fruit set after tagging were
California, Davis, CA 95616 not removed. Fruit were harvested after 3,
Additional index words. Cucumis sativus, parthenocarpic, storage, maturity, growth, 6, 9, 12, 15, 20, 25, and 30 days from tag­
respiration, ethylene, chlorophyll, shelf-life ging.
Individual fruit were enclosed in glass jars
A bstract. Parthenocarpic cucumber fruit (iC ucum is sativus L. cv. Deliva) were har­ of appropriate size to minimize the void space
vested from 3 to 30 days after anthesis. Fresh weight increase followed a single sigmoid and then placed at 20° ± 1°C, under a con­
curve. Commercial maturity was attained 10 to 11 days after anthesis, and these mature tinuous and metered flow of humidified air.
fruit had a 20- to 30-day shelf life and good color retention at 20°C. Chlorophyll content The flow rates were selected to ensure that
at harvest decreased with fruit age. Fruit harvested before commercial maturity showed C 0 2 accumulation did not exceed 0.3%. Un­
declining rates of C 02 and C2H4 production at 20°, while commercially mature fruit less otherwise stated, 4 replications of one
maintained relatively constant rates throughout their holding period. Fruit harvested fruit each were used for each treatment. Car­
after commercial maturity showed increased respiration during holding; this increase bon dioxide was measured daily using an
was greatest for 30-day-old fruit. Fruit harvested 20, 25, and 30 days after anthesis infrared gas analyzer (Horiba PIR-2000) un­
showed peaks of C2H4 production during holding; this production was most pro­ til fruit showed senescent breakdown or de­
nounced for mature fruit. cay. Ethylene was determined using a Carle
Model ACG-211 series S flame ionization
Most studies of cucumber respiration and formly from anthesis to 12 days for pickling gas chromatograph. Visual quality and color
C2H4 production have considered only fruit types and to 14 days for fresh-market types changes were observed every 3 days and as­
harvested at or near table maturity, but of (17). Differences in final fruit length resulted signed as previously described (7). Visual
unspecified age. However, significant phys­ from differences in the period of growth, not quality was rated on a scale of 9 to 1 (9,
iological changes associated with maturation from the rate of growth. Davis and Kempton excellent; 7, good; 5, fair; 3, poor; 1, very
or ripening may occur before or after com­ (3) studied the compositional changes in poor). Color was rated on a scale of 5 to 1
mercial maturity. Previous workers studied greenhouse-grown parthenocarpic cucum­ (5, dark green; 4, green; 3, yellowish green;
respiration, C2H4 production, quality, and bers during growth and maturation. 2, greenish yellow; 1, yellow). Since both
compositional changes of cucumbers under The influence of stage of development at experiments exhibited similar results, only
either different temperature regimes (5, 11) harvest on subsequent respiration and C2H4 the 1983 results are presented.
or modified atmospheres (4, 11). Ethylene production has not been reported for either Samples of peel, about 2 mm thick, were
biosynthesis in cucumbers has been studied seeded or parthenocarpic varieties grown in taken for chlorophyll analysis from the ends
by Wang and Adams (16). Saltveit and greenhouses. This work was conducted to and middle part of 2 individual fruit at each
McFeeters (12) reported C2H4 production rates study the physiology of greenhouse-grown harvest. The samples were frozen at —10°C
and polygalacturonase (PG) activity during parthenocarpic cucumbers harvested at dif­ and subsequently analyzed by the method
development and maturation of pickling cu­ ferent stages of development. In addition, described by Amon (1). The frozen samples
cumbers. The quality of processing cucum­ the relationship between fruit development were homogenized in 80% acetone, centri­
bers has received considerable attention and and chlorophyll content at harvest, between fuged, and chlorophyll content determined
was reviewed by Ennis and O ’Sullivan (6). fruit development and storage life, and color from absorbance at 645, 652, and 663 nm.
Cucumbers have been classified as non­ changes during storage at 20°C were inves­ Growth, as measured by fresh weight (Fig.
climacteric fruit (2). Reportedly they pro­ tigated. 1), followed a single sigmoid curve as ob-
duce little C2H4 during grow th and
development (10, 12), but show a peak in
production after harvest (12). However, res­
piration continued to decrease during the pe­ 40 _
riod of increased C2H4 production, and no E
o
correlation was found between fruit weight
and either the timing or the intensity of C2H4 30 5
production (12). ©
Previous work indicates that the pattern of E
(0
cucumber growth is a single sigmoid curve
20 Q
(3, 17). A photographic analysis of fruit
elongation showed that length increased uni- o
SI
10 "o>
Received for publication 16 Aug. 1985. The cost
of publishing this paper was defrayed in part by
the payment of page charges. Under postal regu­
lations, this paper therefore must be hereby marked
advertisement solely to indicate this fact.
Graduate student. Present address: Department of Days After Anthesis
Horticulture, Univ. of Maryland, College Park, Fig. 1. Growth of ‘D eliva’ cucumbers as measured by fresh weight, length, and diameter o f fruit
MD 20742. harvested at indicated days after anthesis.

HortScience, Vol . 21(5), October 1986 1165

Table 1. Storage life, total life, and chlorophyll content of ‘D eliva’ cucumbers harvested at indicated
days after anthesis.
Fruit age Days after harvest
(days) to reach quality 5Z
3 14 cx
6 31 a
9 31 a
12 22 b
15 13 c
20 11 cd
25 6d
zStorage life is expressed as the number of days after harvest required for the fruit to deteriorate to a
fair condition (score 5).
yStorage was at 20°C. Total life is the number of days from anthesis to a score of 5.
xMean separation by lsd at the 1% level
served by others (3, 17). The rate of weight
increase was almost linear from 3 to 20 days.
Fruit continued to gain weight and increase
in length up to 30 days, but at reduced rates.
In contrast, previous studies found that in­
crease in length almost ceased after 21 days
(3) or 14 days (17). Fruit diameter increased
up to 20 days and then remained fairly con­
stant. During the summer, fruit reached the
common commercial harvest weight (0.5
kg)— about half of the final weight—by the
11th day.
After an initial decrease, chlorophyll con­
tent of the peel remained fairly constant from
6 to 12 days after anthesis and then de­
creased uniformly as the fruit developed on
the plant (Table 1). Takama et al. (13) found
a constant decrease in chlorophyll content,
but they only measured changes during the
first 10 days of development.
Respiratory patterns for fruit harvested up
to 25 days from anthesis (Fig. 2) were gen­
erally similar to those reported for other non­
climacteric fruit and other kinds of cucumbers
(5, 10). The initial and subsequent respira­
tion rates of younger fruit (3 and 6 days old)
were high compared to older fruit (9 to 20
days old), which produced C 0 2 at a rela­
tively constant rate throughout the post­
harvest period. Respiration of 30-day-old fruit
showed a statistically significant (5% level)
1.6-fold increase after the first day before
decreasing 3 days later. Only fruit harvested
at this age showed a significant increase in
respiration. In retrospect, it is evident that
fruit of more-advanced development should
have been included in the study.
Rates of C2H4 production followed 3 in­
teresting patterns (Fig. 3). First, there was a
marked decline with time in C2H4 produc­
tion by fruit harvested 3 days after anthesis.
Second, 6- to 15-day-old fruit showed fairly
constant rates until senescence occurred.
Third, 20- to 30-day-old fruit showed a tran­
sient 3- to 5-fold increase in C2H4 produc­
tion after harvest, which was greater for the
more-mature fruit. Although the variation
among individual fruit within each harvest
was great, the trend was similar. The in­
crease was significant at the 5% level only
for 25- and 30-day-old fruit.
Saltveit and McFeeters (12) found that cu­
cumbers of various stages of development
produced a transitory rise in C2H4 produc­
tion from 6 to 25 days after harvest, although
1166
Days after anthesis Chlorophyll
to reach quality 5y (m g -g -1)
17 0.92
37 0.54
40 0.53
34 0.52
28 0.32
31 0.21
31 0.10
C 0 2 production declined constantly during
this time. In addition, this transitory rise in
C2H4 production was of reduced intensity as Days After Anthesis
the time of its occurrence after harvest in­ Fig. 2. Respiration rates (C 0 2, mg*kg- 1*hr-1)
creased. for cucumbers harvested at indicated days after
Climacteric fruit such as melons and to­ anthesis and held at 20°C. The values represent
matoes tend to show an increase in C2H4 and the averages of 4 fruit, except 3 in the case of
30-day-old fruit. lsd0.05 - 4.5 for 30-day-old
C 0 2 production and final senescence at about
fruit. Increases in respiration rates by 25-day-
the same ultimate age after anthesis, regard­ old fruit were not significant at the 5% level.
less of the age at harvest (8). In this study,
however, mature cucumbers tended to show
an increase in C 0 2 and C2H4 production rates
at the same time after harvest and not at the
same time after anthesis (Figs. 2 and 3). A
possible explanation is that as the fruit de­
velop, they have an increased potential to
produce C2H4 and C 0 2, and this production
is induced by harvest. This increased poten­
tial may be related to a decrease in inhibitory
factors in maturing fruit, which permits the
expression of C2H4 producing capabilities,
or an increase in C2H4 producing capabili­
ties, or a combination of both.
Loss of green color and firmness were im­
portant symptoms of deterioration and were
first noted at the stem end. Pathological
breakdown also appeared first at the stem
end. All fruit developed raised spots or
“ blisters” of darker green color. Yellow color
developed around the blisters before they be­
came yellow, giving the fruit a mosaic-like Days After Anthesis
appearance. Blisters also developed on the Fig. 3. Ethylene production rates (nl*kg_ 1*hr-1)
mature fruit while they were still attached to for cucumbers harvested at indicated days after
the vine. Robinson et al. (11) indicated that anthesis and held at 20°C. The values represent
the averages of 4 fruit, except 3 in the case of
blistering is one of the storage disorders of
30-day-old fruit. lsd0 05 = 85.0 for 30-day-old
cucumbers. Papadakis (9) reported that these
fruit and 49.7 for 25-day-old fruit. Increases in
raised spots appeared on all fruit held at 12.5° C 2H4 production rates by 20-day-old fruit were
and 15°C but not on fruit at 5°. Examination not significant at the 5% level.
of the spots under a light microscope did not
reveal bacteria or fungi. A similar physio­
logical disorder (called tumor or waxy blis­ shelf life. However, for maximum shelf life,
ter) appears on m ature-green tomatoes. it appears desirable to harvest somewhat ear­
Treshow (15) studied the tomato tumors and lier within the acceptable size range.
concluded that rubbing encountered during Total life from anthesis to quality 5 was
picking and handling of green fruit was the reasonably constant for all fruit that were
main cause of this disorder. harvested 12 or more days after anthesis (Ta­
Storage life was expressed as the days re­ ble 1). Thus, shelf life after harvest is re­
quired to deteriorate to a quality score of 5. duced for the advanced stages of development.
Fruit harvested 6 and 9 days after anthesis Patterns of quality deterioration and color
had a shelf life of 31 days, significantly longer change were similar for the 3 harvests made
than that for all other ages (Table 1). The within the range of usual commercial prac­
12-day-old fruit had a shelf life of 22 days. tice (i.e., 9-, 12-, and 15-day harvests).
Since the usual harvest size of 0.5 kg was Changes were relatively slow during the early
attained at 11 days, it is concluded that fruit part of the holding period and then acceler­
of commercial maturity have a relatively long ated.
HortScience, V ol . 21(5), October 1986
 Based on the results reported here and by
others, it is concluded that cucumbers of
commercial table maturity (9 to 15 days) be­
have like nonclimacteric fruit. However, since
most mature fruit do produce C2H4 and show
an increasing rate of C 0 2 production, more
research is needed to establish whether or not
ripening cucumbers show a climacteric re­
sponse. Such work should include fruit that
have developed for 30 days and more on the
plant. Carbon dioxide and C2H4 production
should be determined for both attached and
harvested fruit. Furthermore, their responses
to different levels of propylene would help
establish their classification as climacteric or
nonclimacteric fruit.
1.
Literature Cited
Amon, D.I. 1949. Copper enzymes in iso­
lated chloroplasts. Polyphenoloxidase in Beta
vulgaris. Plant Physiol. 24:1-15.
2. Biale, J.B. and R.E. Young. 1981. Respi­
ration and ripening in fruit— retrospect and
prospect, p. 1-39. In J. Friend and M.J.C.
Rhodes (eds.). Recent advances in the bio­
chem istry o f fruit and vegetab les. A ca­
demic, New York.
3. Davis, J.N. and R.J. Kempton. 1976. Some
changes in the composition o f the fruit of
the glasshouse cucumber (Cucumis sativus
L.) during growth, maturation and senes­
cence. J. Sci. Food Agr. 27:413-418.
4. Eaks, I.L. 1956. Effect of modified atmos­
pheres on cucumbers at chilling and non­
chilling temperatures. Proc. Amer. Soc. Hort.
Sci. 67:473-478.
5. Eaks, I.L. and L.L. Morris. 1956. Respi­
ration of cucumber fruit associated with
physiological injury at chilling tempera­
tures. Plant Physiol. 31:308-314.
6. Ennis, D.M . and J. O ’Sullivan. 1979. Cuc­
umber qu ality— a rev iew . J. Food S ci.
44:186-189.
7. K anellis, A .K . 1984. P hysiological re­
sponses of cucumber fruit to stage of de­
velopment at harvest and to oxygen levels
in the storage atmosphere. MS Thesis, Univ.
of California, Davis.
8. Mizuno, S. and H.K. Pratt. 1973. Relations
of repsiration and ethylene production to
maturity in the watermelon. J. Amer. Soc.
Hort. Sci. 98:614—617.
9. Papadakis, C.M. 1981. Effect of nitrogen
and temperature on the performance of Eu­
ropean cucumber, Cucumis sativus L ., using
soil-less culture. MS Thesis, Univ. o f Cal­
ifornia, Davis.
10. Poenicke, E .F ., S.J. Kays, D .A . Smittle,
and R.E. Williamson. 1977. Ethylene in re­
lation to postharvest quality deterioration in
processing cucumbers. J. Amer. Soc. Hort.
Sci. 102:303-306.
11. Robinson, J.E., K.M. Browne, and W.G.
Burton. 1975. Storage characteristics of some
vegetable and soft fruit. Ann. Applied Biol.
81:399-408.
12. Saltveit, M.E. Jr., and R.F. McFeeters. 1980.
Polygalacturonase activity and ethylene syn­
thesis during cucumber fruit development and
maturation. Plant Physiol. 66:1019-1023.
13. Takama, R ., S. Fukuda, Y. Toyomaki, K.
Toyomaki, and S. Saito. 1973. Changes of
chemical components of cucumber and sweet
pepper fruit during growth on the tree. J.
Jpn. Soc. Food & Nutr. 26:329.
14. Tiedjens, V .A . 1928. The relation of envi­
HortScience, Vol . 21(5), October 1986
ronment to shape of fruit in Cucumis stivus ing-induced ethylene production in cucum­
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of the plants. J. Agr. Res. 36:795-809. 69:424-442.
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HortScience 21(5): 1167-1169. 1986.
Controlled Atmosphere Effects on the
Pathogenicity of Fungi on Celery and
on the Growth of Botrytis cinerea
Andres A. Reyes and Richard B. Smith
Agriculture Canada Research Station and Horticultural Research Institute
of Ontario, Vineland Station, Ontario, Canada LOR 2E0
Additional index words. Apium graveolens, postharvest diseases, cold storage
A bstract. Sclerotin ia sclerotioru m (Lib.) de Bary and B o trytis cin erea Pers. were highly
pathogenic to celery stored at 0° to 1°C in normal air (21% 0 2). A ltern a ria d au ci (Kuhn)
Groves & Skolko, R h izopu s nigrican s Ehrenb., P en icilliu m sp., and F u sariu m ox y­
sporum Schlecht. were nonpathogenic. An atmosphere of 7.5% CO/1.5% 0 2 was more
suppressive to disease caused by B . cin erea and S. sclerotioru m than low 1.5% 0 2
atmosphere alone. The 4% C 0 2/1.5% 0 2 and 0.0003% C2H4/1.5% 0 2 atmospheres
were slightly suppressive to disease caused by S. sclerotioru m only. The 7.5% CO/1.5%
0 2 atmosphere also was consistently suppressive to mycelial growth, spore germination,
and germ tube elongation of B . cin erea .
Vegetables have the potential to retain better phal tip cultures of S. sclerotiorum, B. ci­
marketable quality when stored in controlled nerea, A. dauci, R. nigricans, Pencillium
atmosphere (CA) than when kept in normal sp., and F. oxysporum. They were main­
air (6, 11). This improved marketable qual­ tained on potato dextrose agar (PDA) plates
ity may be due partly to the suppression of (100 x 15 mm) at 21° ± 1°C. Eight-mil­
postharvest diseases (1, 8, 10). On the other limeter-diameter agar plugs were prepared
hand, C2H4 in the storage atmosphere has from the margin of 5-day-old cultures. A spore
been reported to stimulate postharvest decay suspension (8 X 106 spores per milliliter) of
(4, 5). B. cinerea was prepared aseptically from 3-
In a preliminary study, species of Alter- week-old PDA plate cultures maintained at
naria, Botrytis, Fusarium, Penicillium, Rhi- 21° ± 1°.
zopus, and Sclerotinia were isolated from Storage atmospheres consisted of 1.5% 0 2
celery (Apium graveolens L. var. dulce DC.) either alone (low 0 2) or with 0.0003% (3
after 8 weeks of storage at 0° to 1°C in nor­ ppm) ethylene (C2H4), 4% C 0 2, or 7.5%
mal air (unpublished data). The present study CO, the remainder being N. These atmo­
was initiated to determine the effects of CA spheres were prepared by Union Carbide
on the severity of diseases caused by these Canada in 60-hl pressure cylinders. The con­
fungi on celery and on the in vitro growth trol (air) contained 21% 0 2.
of B. cinerea, a common storage pathogen Fresh field celery plants (‘Utah 52-70’)
of agricultural crops. were trimmed to a 15-cm length, placed up­
The fungi used were single spore or hy- right in 48 x 24 x 17 cm plastic boxes (10
plants per box), immersed for 60 sec in 0.6%
NaOCl solution, and rinsed 3 times with sterile
distilled water.
Received for publication 15 Oct. 1985. We thank
Each of 3 randomly selected petioles per
Ann M. Curwin and Sharon E. Stevenson for tech­
nical help. The cost of publishing this paper was
plant was inoculated aseptically by placing
defrayed in part by the payment of page charges. an agar plug of inoculum on the cut end. An
Under postal regulations, this paper therefore must agar plug without fungus served as the con­
be hereby marked advertisement solely to indicate trol. Two plants were used for each fungus,
this fact. and the plants were separated with plastic
1167