J. Amer. Soc. Hort. Sci. 104(6):835—838. 1979.

Fruit Set and Development of Seeded and Seedless Tomato

Fruits under Diverse Regimes of Temperature and
Pollination1
Irena Rylski
Institute o f Field and Garden Crops, The Volcani Center, Bet Dagan, Israel
Additional index words. Lycopersicon esculentum
Abstract. Fruit set and fruit development of seeded, naturally seedless, and growth-regulator-induced seedless
tomatoes (Lycopersicon esculentum Mill. cv. Azes), were examined under 17°, 22° and 27°C day temperatures,
all with natural daylight and constant 10° night temperature. The best seeded fruit set and development were
at 22° day temperature. A high positive correlation (r = 0.9) was found between number of seeds per fruit and
fruit size with a day temperature of 27°; there was a lower but still positive correlation with lower temperature.
Seedless fruit developed, and attained marketable weight (above 40 g), only when the inflorescences had been
sprayed with 20% ft naphthoxyacetic acid (B-NOA). Day temperature also affected fruit shape. At 27° the fruit
was almost globe-shaped, but at the lower temperatures the fruit became more oblate. Fruit puffiness was par-
ticularly related to lack of fertilization resulting from flower emasculation or to lower day temperature affecting
set of seed-deficient fruits. “No-Seed” treatment led to green jelly production and pointed blossom-end in fruits,
under all temperature regimes.

Polyethylene (PE)-covered tunnels, and net, polyethylene -
or glass-covered greenhouses are used in Israel for winter-grown
tomatoes, without additional heating. Under these conditions
the night temperature may fall below 10°C, approximately
equaling the outside temperature. However, day temperatures
vary considerably throughout the winter growing season, as
a result of different levels of radiation, type of covering, rate of
ventilation, and other factors.
Lack of tomato fruit set, which is common under these
particular conditions, can be overcome by the use of growth
regulators. This has been investigated in Israel in both unpro-
tected (9) and protected (5, 11, 13) tomato crops and become
accepted agricultural practice. However, sometimes many small,
seedless or seed deficient fruits are produced.
Since under protected growing conditions it is possible to
regulate day temperatures to some extent by appropriate
cover and ventilation, an experiment was performed under con-
trolled temperatures to investigate the effect of day tempera-
tures under low night temperature conditions on fruit set and
fruit development in both seeded and seedless fruits, produced
either naturally or as a result of growth-regulator treatment.
Materials and Methods
The experiment was performed with indeterminate ‘Azes’
tomato plants pruned to 1 branch, the usual practice in green-
houses. Plants at the 6-leaf stage, grown in a greenhouse in
10-liter plastic containers containing a mixture of 1 loam soil: 1
peat:l vermiculite (by volume) at the rate of were transferred
to 3 glass-covered controlled growth chambers which permit
plant growth under natural light conditions.
The day temperatures tested were 17°, 22° and 27°C (all
± 1°) for 12 hr; night temperatures were the same for all treat-
ments, viz., 10° (± 1°). There were 16 plants completely
randomized (once a week the containers were moved to a new
site) in each chamber and these were subjected to 4 different
treatments: 1) open pollination (control); 2) non-pollinated
(emasculated) flowers; 3) hand-pollinated (emasculated and
pollinated flowers); and 4) open-pollinated flowers, each inflo-
1Received for publication June 13, 1978. Contribution from the Agri-
cultural Research Organization, The Volcani Center, Bet Dagan, Israel.
No. 151-E, 1978 series.
The cost of publishing this paper was defrayed in part by the payment of
page charges. Under postal regulations, this paper must therefore be
hereby marked advertisement solely to indicate this fact.
rescence sprayed twice with ca. 0.5cc of a 0.5% solution of
20% B-NOA (“No-Seed”). The first spray treatment was given
when there were three open flowers on the inflorescence, and
the second one was given 1 week later. In treatments 2 and 3
the flowers were emasculated 1 day prior to anthesis. In treat-
ment 3, pollination was performed the day after emasculation.
The pollen was collected from flowers of the same cultivar
grown in a greenhouse in which temperatures were not allowed
to drop below 14°. On 5 inflorescences, numbers 2-6 per plant,
all flowers were tagged at anthesis and fruit set was determined.
Each fruit was picked individually when turning red and
examined for weight, shape (length/diameter ratio), number
of seeds, puffiness, and color of the jelly. Puffiness was rated
according to separation of the pericarp from the endocarp,
as follows: slightly puffy — no more than % of pericarp sepa-
rated, moderately puffy —% to xh separated, very puffy - more
than Vi separated. Jelly was rated visually as red, pale green or
dark green.
Results
Flowering. Growing tomato plants under different day
regimes, viz., 17°, 22° and 27°C, brought about changes in the
rate of development so that their first 5 inflorescences reached
anthesis at different periods, viz., 126, 100 and 92 days after
sowing respectively. However, with rising temperature there
was a significant drop in the average number of flowers per
inflorescence: 13.1, 11.0 and 9.4 flowers at 17°, 22° and 27°,
respectively.
Fruit set. The highest percent of seeded fruit set was ob-
tained under the 22° temperature regime, for all inflorescence
treatments (Table 1). At this temperature, with the open-
and hand-pollinated treatments, the percentage of seeded
fruit was high and almost identical to total fruit set (seeded
plus non-seeded fruit). At 27°, all fruits which set contained
seeds, but fruit set was lower than at 22°. At 17°, with open-
pollination, a high percentage of the flowers set fruit but only
about half of them were seeded. Hand pollination increased the
rate of seeded fruit set at 17° and 22°, at which temperatures
there was little abortion of emasculated and non-pollinated
flowers. However, at 27°, flower abortion was severe. B-NOA
treatment increased the rate of seedless fruit set.
Fruit size. Seedless fruit attained marketable weight (above
40 g) only in those cases in which the inflorescence had been
sprayed with B-NOA at 22° or 17°C. We observed 23% of the
total number of fruit was seedless and marketable at 22° and
49% at 17°. At 22°, 80% of the fruits in all treatments, except

J. Amer. Soc. Hort. Sci. 104(6):835-838. 1979. 835

 Table 1. Effect of day temperature on total and seeded fruit-set after different flower treatments.

Non-pollinated, Fruit set (%)

Day
emasculated Open-pollinated Hand-pollinated B-NOA
temp
(°C) Total2 Seeded Total Seeded Total Seeded Total Seeded

17 91aAy 0 93aA 53bB 85 aA 85 aA 84 aA 33aB

22 94aA 0 81aAB 78aA 91aA 91aA 71bB 51aB
27 19bB 0 5 9b A 5 8 bA 50bA 5 0b A 6 2b A 46aA

including fruits with and without seeds.

yMean separation within columns by Duncan’s multiple range test, 5% level. Lower case letters refer to temperature treat-
ments, upper case letters to inflorescence treatments.

those developing from emasculated flowers (non-pollinated parthenocarpic fruits (non-pollinated), jelly development it-
and no auxin treatment), were above 40 g. However seedless self was very restricted and consequently the percentage of
fruits were smaller, on the average, than seeded ones; about 50% these fruits with green jelly was limited.
of the seeded fruits weighed over 60 g, whereas only 35% of
the seedless fruit attained this weight. Moreover, at 17°C, 60% Discussion
of all the fruits which set as a result of open pollination weighed The decline in number of flowers per inflorescence as tem-
less than 40 g, and in the seedless fruit of this treatment, 95% peratures rose, is a well known occurrence in tomatoes (1, 6).
weighed less than 40 g. B-NOA treatment increased fruit size, In the present work, plants were grown from the 6-leaf stage,
including that of seedless fruit. In the 27° treatment, fruit that is, after differentiation of the first inflorescences (1),
weight was lower than in the other regimes, for all inflores- under low night temperature conditions (10°C), and various day
cence treatments. With B-NOA treatments at 27°C, fruit tended temperatures (17°, 22° and 27°C). Nevertheless, the number of
to be somewhat smaller than that from non-treated flowers. flowers per inflorescence, from the 2nd to the 5th, declined
Number o f seeds per fruit and fruit size. At 17°C most of with rising day temperature. Of all the environmental condi-
the fruits contained fewer than 100 seeds per fruit (Table 2), tions, temperature has undoubtedly the most significant effect
but at 22° and 27°C more than 50% contained over 100 seeds. on fruit set in tomatoes. Went (18) stressed this in greater
There was a high (r = 0.9) correlation, calculated from all fruits detail when he established that it is the night temperature
obtained from open- and hand-pollinated flowers, between
number of seeds per fruit and fruit size, the coefficient increas-
ing with rising temperature (Fig. 1).
Number o f days from anthesis until fruit ripening. This
variable decreased from 78 days at 17°C to 70 days at 22°
and 50 days at 27°. No relationship was found between num-
ber of seeds per fruit and number of days from anthesis to
ripening.
Fruit quality. At high day temperature (27°C) fruit was
almost round, whereas at 17° fruit was more oblate (Table 3).
Non-fertilized small fruits which developed under low tempera-
ture, were fasciated. A somewhat pointed blossom-end was
found in all B-NOA-treated fruit, at all temperatures. At 17°
a high percentage of the seedless fruits were puffy (Fig. 2).
At this temperature about one-third of the fruits produced as
a result of open pollination were puffy. With the rise in temper-
ature, the percentage of puffy fruit declined and at 27° puffi-
ness was found only in parthenocarpic fruit. Green jelly was
particularly evident in B-NOA-treated fruit, under all tempera-
ture regimes. At 17°, about one-third of the fruits from open-
and hand-pollination, contained green jelly. In small, puffy,

Table 2. Fruit distribution according to seeds per

fruit (% of total no. of fruits) at different day
temperatures.

Fruit distribution (%)Z

Day temperature
Seeds/fruit 17°C 22° 27°

1-20 30 8 8
21-100 51 25 36
> 1 0 0 19 67 56 Fig. 1. Relationship between number of seeds/fruit and fruit weight
under different day time temperatures. Fruits obtained from open-
zDifferences between groups of fruits and tempera- pollinated and hand-pollinated flowers. Numbers beside dots show
ture conditions significant at 1 % level. number of fruits examined.

836 J. Amer. Soc. Hort. Sci. 104(6):835-838. 1979.

 Table 3. Effect of day temperature on fruit shape from different flower treatments.

Fruit shape
Slightly Moderately
pointed pointed
Day No. Length/ blossom blossom
temp fruits diameter end end Fasciated
(°C) Treatment examined ratio (%) (%) (%)

17 Non-pollinated emasculated 173 0.87 0 0 58

Open-pollinated 112 0.87 8 1 14
Hand-pollinated 153 0 .8 6 3 0 0

B-NOA 174 0 .8 6 23 10 1

22 Non-pollinated emasculated 171 0.87 0 0 1

Open-pollinated 98 0.93 0 0 0

Hand-pollinated 161 0 .8 8 1 0 0

B-NOA 134 0.89 22 4 0

27 Non-pollinated emasculated 13 0 0 0

Open-pollinated 61 0.95 0 0 0

Hand-pollinated 89 0.95 0 0 0

B-NOA 93 0.96 22 2 0

27 °C 22 °C 17 °C
OPEN POLLINATED which determines fruit set, with 15-20° being optimal. Accord-
ing to Went (18), at temperatures above or below this range, the
rate of fruit set declines, down to complete lack of fruit set.
Robinson et al. (10) demonstrated that in tomatoes, low
temperatures have an adverse effect on fruit set, especially in
their influence on gametogenesis, in particular on the produc-
tion of pollen grains. In experiments performed by Robinson
et al. (10) similar to those described by Charles & Harris (2),
a temperature of 10° at the time of meiosis brought about
production of nonviable pollen grains. Consequently, Robinson
et al. (10), as Went (17), suggested that temperature should not
be allowed to drop below 16°C during flowering, so as to
ensure regular fruit set. However, in the present experiment,
there was a high rate of seeded fruit set even under low night
temperature conditions (10° ± 1°), provided that the day
temperature was 22°. Our experiment was performed under
HAND POLLINATED natural light conditions. This leads to the conclusion that other
environmental effects must also be taken into account when
considering temperature. These include the combination of day
and night temperature, light intensity, air humidity, and geno-
type. Schaible (14) obtained fruit set in tomatoes at a night
temperature declared maximal by Went (17), and Curme (3)
obtained satisfactory fruit set in several cultivars with a night
temperature of 7°. In an experiment performed in summer,
when light intensity was greater, Curme (3) obtained better
fruit set than in a similar experiment performed in winter,
under the same temperature conditions.
When pollination and fertilization do not take place, the
flowers abort, or, under specific environmental conditions,
there is parthenocarpic fruit set. In the present experiment,
under high day (27°C) and low night (10°) temperatures,
about 80% of non-fertilized flowers aborted. Under the same
night temperature but with low day temperature (17°), non-
PUFFINESS fertilized flowers set parthenocparic fruits, but these remained
NOT PUFFY small until ripening, unless treated with growth regulators.
Treatments with appropriate growth regulators induce fruit
SLIGHTLY PUFFY growth both in fertilized and in non-fertilized fruit. These may
limit seed production in fruits, according to Verkerk (16), as
MODERATELY PUFFY a result of the abundant flow of auxin to the ovules before
fertilization, which prevents subsequent fertilization. H ow ever,
VERY PUFFY ovules are likely to abort after fertilization, as in the present
experiment where flowers which reached anthesis several days
Fig. 2. The rate of fruit puffiness (right) and production of green jelly before the B-NOA treatment nevertheless produced seed-
(left) under different day temperature conditions. Each half circle deficient fruits. Where the B-NOA treatments brought about
represents 1 0 0 % of the fruit examined (number of fruits as in table 2 ).

J. Amer. Soc. Hort. Sci. 104(6):835-838. 1979. 837

increased fruit size under the 17° day temperature regime,
fruit size was reduced under the 27° regime. From this we
conclude that under protected growing conditions, appropriate
ventilation during the daytime is essential to control and main-
tain the ambient air temperature at a level optimal to growth-
regulator-treated, as well as pollinated, fruit growth.
Various experiments (4, 15, 16) have shown that the final
size of the tomato fruit depends on the number of seeds de-
veloping within it. Varga and Bruinsma (15) qualify this condi-
tion to a minimum of 9 seeds.
In the present work it was found that even a few seeds can
bring about a certain amount of fruit growth under low temper-
ature conditions (10°C by night and 17° by day). At these
temperatures even non-fertilized ovaries do not drop. There
was a high positive correlation (r=0.9) between number of
seeds per fruit and fruit size at 27°, a temperature at which only
fertilized fruit can develop. At lower temperatures, which tend
to promote parthenocarpic fruit set, the correlation coefficient
between number of seeds per fruit and fruit size was lower.
In this work no relationship was found between the number
of seeds per fruit and the number of days required until fruit
ripening, after anthesis, in contrast to what may have been
expected. Nitsch et al. (8) examining the physiological role of
the seeds in fruit development, found that the amount of endog-
enous auxins in the fruit increases after fertilization, and the
formation of the endosperm and embryo induces cell enlarge-
ment in the fruit.
Marre and Murneek (7) consider seeds to be the center of
auxin production, which in turn controls the metabolism of
carbohydrates. Consequently, fruits with many seeds may be
endowed with heightened metabolic activity, accelerating cell
enlargement and fruit ripening. Varga and Bruinsma (15) con-
cluded that fruit growth in tomatoes should be attributed to
the sink activity of the seeds and not to cell enlargement as
a result of auxin production in the course of seed develop-
ment. In our work we did not find that a larger number of
seeds per fruit accelerated fruit ripening, whereas Verkerk
(16), found a reduction in the period from anthesis to fruit
ripening by 4-7 days.
Apparently, both temperature and auxin affect fruit quality,
as judged by shape, puffiness and jelly color. Low night and/or
day temperature at the early stage of flower development causes
changes in ovary structure and, consequently, in fruit shape.
Different cultivars of tomatoes differ in their sensitivity to low
temperature and auxin treatments, as well as in their suscepti-
838
bility to fruit puffiness (12). In 4Azes’, the rate of puffiness was
related to low temperature and lack of seed production. The
green jelly production within the fruit was related to the appli-
cation of B-NOA and to low day temperature.
Literature Cited
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and low temperatures. Can. J. Plant Sci. 52:497-506.
3. Curme, J. H. 1962. Effect of low night temperatures on tomato
fruit set. Proc. Plant Sci. Symp., Campbell Soup Co., Camden,
N.J. p. 89-98.
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J. Amer. Soc. Hort. Sci. 104(6):835—838 1979.