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Khokhar, K.M., 2008. Effect of temperature and photoperiod on the incidence

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Journal of Horticultural Science & Biotechnology (2008) 83 (4) 488–496

Effect of temperature and photoperiod on the incidence of bulbing

and bolting in seedlings of onion cultivars of diverse origin

By KHALID MAHMUD KHOKHAR

Vegetable Crops Research Programme, Horticulture Research Institute, National Agricultural
Research Centre, Park Road, Islamabad 44000, Pakistan
(e-mail: Tokmahmud@yahoo.com) (Accepted 10 January 2008)

SUMMARY
The effects of temperature and photoperiod on bulbing and bolting in onion (Allium cepa L.) cultivars: ‘Senshyu
Yellow’, ‘Jaune Espagnol’, ‘Hygro’, ‘Sito’, ‘Delta’, ‘Australian Brown’, ‘Rijnsburger Balstora’, and ‘Phulkara’ from
Japan, Turkey, The Netherlands, The United Kingdom, Australia, and Pakistan were examined under controlled
environments in glasshouse conditions in the UK. Plants of each cultivar, at the three-leaf stage, were transferred to a
wide range of photo-thermal regimes consisting of six-set point temperatures (6°, 10°, 14°, 18°, 22° or 26°C) and four
photoperiods (8, 11, 14 or 17 h d–1). The bulbing ratio in all cultivars increased curvi-linearly with increasing
temperature and lengthening photoperiod. Increasing photoperiod promoted bulbing, while under very short
photoperiods (8 h d–1) no cultivars bulbed, even after 60 d of growth. The time to bulb maturity decreased linearly with
increasing temperature and lengthening photoperiod. Under low temperatures, the time to floral initiation shortened
with lengthening photoperiods (8 – 14 h d–1). Under photo-thermal regimes favouring floral initiation, leaf number
increased with both increasing temperature and lengthening photoperiod.

I t is well-established that long-days (LD) and high

temperatures promote onion bulbing (Heath, 1945;
Kato, 1964). If day-length is sufficiently short, no bulbing
occurs, even at high temperatures (e.g., 34°C; Heath,
1943, 1945; Kato, 1964; Butt, 1968; Steer, 1980; Terabun,
1980), while bulbing is delayed or prevented by low
temperatures (Heath, 1943; 1945).
In contrast, a period of low temperatures, between
7° – 13°C for 20 – 90 d, leads to flower induction in most
onion cultivars (Shishido and Saito, 1975; 1977; Brewster,
1983; Bertaud, 1988; De Bon and Rhino, 1988; Peters,
1990). Extended photoperiods under low temperatures
have been shown to promote the initiation of
inflorescences (Woodbury, 1950; Shishido and Saito,
1975; Brewster, 1983). However, Holdsworth and Heath
(1950) and van Kampen (1970) reported that
photoperiod had no effect on the initiation of
inflorescences. Brewster (1987b) showed that the rate of
progress to flower initiation was more-or-less a linear
function of photoperiod between 8 – 20 h d–1 under
inducing temperatures. Studies on both bulbs and
seedlings have shown that flower initiation starts only
after a juvenile phase, by which time the plant has a
minimum number of four-to-14 leaves, including leaf
initials (Heath and Mathur, 1944; Hartsema, 1947; Heath
and Holdsworth, 1948; Ito, 1956; Shishido and Saito, 1976;
Rabinowitch, 1985; 1990).
Once an inflorescence has been initiated, its rate of
development increases with increases in temperature
(10° – 30°C) and lengthening of the photoperiod (14 – 16
h d–1; Heath and Mathur, 1944; Heath, 1945; Woodbury,
1950; Brewster, 1982a; 1983; 1987a; Bertaud, 1988).
However, high temperatures (e.g., 30°C for cv.
‘Rijnsburger’) during early development of the scape can
result in flower abortion because of competition for
assimilates from bulbing, which may be induced if the
day-length is appropriate for the cultivar (van Kampen,
1970).
When breeding onions, the interval between one
generation and the next needs to be minimised.
Normally, a 2 year seed-to-seed cycle occurs, with 1 year
needed to produce the bulbs which then produce seed in
the following year (van Kampen, 1970; Pike, 1986;
Bertaud, 1988; Dowker, 1990). We need to understand
how to achieve an early induction of flowering, for rapid
seed production in breeding programmes, and how to
prevent bolting in seed-sown, over-wintered crops. There
have been limited studies on the effects of temperature
and photoperiod on flowering of onions during their
growing phase, using only a narrow range of
environmental conditions (Shishido and Saito, 1975).
However, some work has been reported on the effect of
temperature on onion plants (Davis and Jones, 1944;
Shishido and Saito, 1976; 1977).
The present studies were carried out to examine the
quantitative effects of a wide range of photo-thermal
conditions on the initiation of inflorescences and bulb
development in a diverse range of onion genotypes, with
the aim of defining the optimum environment for rapid
seed production and for bulbing. Information on an
extended range of post-planting environments would
also be useful for planning a seed production
programme. A seed-to-seed production method would
help to shorten the onion seed production cycle.
 K. M. KHOKHAR
MATERIALS AND METHODS
Seeds of eight onion (Allium cepa L.) cultivars:
‘Senshyu Yellow’, ‘Jaune Espagnol’, ‘Hygro’, ‘Sito’,
‘Delta’, ‘Australian Brown’, ‘Rijnsburger Balstora’ and
‘Phulkara’, originating in Japan, Turkey, the Netherlands,
the United Kingdom, Australia, and Pakistan (Table I)
were sown on 15 April 1996 in P 80 plug trays (TEKU
Plug Trays; BHGS Ltd., Evesham, UK) where each cell
had a diameter of 3 cm, using a peat-based seed-sowing
modular compost (SHL; William Sinclair Horticulture
and Leisure Limited, Lincoln, UK), mixed thoroughly
with 20% (v/v) perlite. After filling the trays, the compost
was firmed lightly then levelled-off. Sangral 111 (William
Sinclair Horticulture and Leisure Limited) was applied
as a supplement, as liquid feed at each watering with an
EC of 1.5 dS m–1 (13.0 mM N; 2.50 mM P; 3.80 mM K),
and a pH of 5.8. The trays were immediately placed in a
glasshouse where the temperature averaged 22.3°C. One
month after sowing, when seedlings had reached the
three-leaf stage, plants were re-potted into 7.5 cm-
diameter pots using a potting medium consisting of
80% (v/v) Fisons Levington M2 peat-based compost, and
20% (v/v) medium-size Silvaperl perlite (William
Sinclair Horticulture Limited).
A factorial combination of eight cultivars, four
photoperiods and six temperature treatments gave a
total of 192 treatment combinations, in each of which
were nine plants. To study the combined effects of
temperature and photoperiod on the bulbing and
flowering responses of eight onion cultivars, plants were
placed on moveable trolleys and exposed (23 May 1996)
to 24 different photo-thermal regimes created by a
factorial combination of six temperatures and four
photoperiods. Sixteen additional plants of each cultivar
were also placed in each treatment combination for
dissection, in order to observe inflorescence initials. Two
plants of each cultivar were selected at random from the
extra plants and dissected 12 weeks after shifting the
plants to the photo-thermal regimes, and thereafter at
14 d intervals until the end of the experiment.
Controlled glasshouse compartments (3.7 m
7 m)
provided minimum temperatures of 6°, 10°, 14°, 18°, 22°,
or 26°C. Each compartment was equipped with four
controlled photoperiod chambers, sealed from exterior
light sources. Day-lengths were extended inside each
chamber by low irradiance lighting (PAR 11 µmol m–2
s–1) provided by one 40 W tungsten plus one 15 W
compact fluorescent light bulb, to provide 8, 11, 14 or 17
h d–1. All trolleys were removed from the photoperiod
chambers at 08.00 h so that plants were exposed to 8 h of
natural daylight in the glasshouse before being returned
to the photoperiod chambers at 16.00 h each day. In all
treatments, the lamps were switched on automatically at
16.00 h to provide day-lengths of 8, 11, 14 or 17 h d–1.
TABLE I
Details of onion cultivars used in this study
Name of cultivar Source
‘Hygro’ Moles Seeds, UK
‘Delta’ Moles Seeds, UK
‘Senshyu Yellow’ A.L. Tozer Ltd, UK
‘Jaune Espagnol’ Genetic Resources Unit, HRI, Wellsbourne, UK
‘Sito’ Nickerson Seeds Ltd, UK
‘Australian Brown’ Genetic Resources Unit, HRI, Wellsbourne, UK
‘Rijnsburger Balstora’ A.L. Tozer Ltd, UK
‘Phulkara’ NARC, Pakistan
489
Inside the chambers, plants were ventilated continuously
at an average air speed of 0.2 m s–1 to maintain
temperature. Actual mean diurnal temperatures within
each compartment were calculated from the
temperatures recorded on a data-logger (Datataker, DT
500; Data Electronics, Letchworth Garden City, UK),
linked to aspirated PT 100 temperature sensors (15 s
scans, logged each hour). The actual mean temperature
for each respective developmental stage (i.e., bulbing,
neckfall, floral initiation) for each photo-thermal
combination was calculated. Plants were sprayed with 0.8
g l–1 BenlateTM (DuPont, Wilmington, DE, USA) 6 weeks
after planting as a preventive measure against fungal
diseases. This was repeated every 14 d until the end of the
experiment. The plants were watered three-times a week,
or more frequently when required. Observations on
bulbing ratios, times to floral initiation, times to bulb
maturation, and leaf numbers were recorded. Bulbing
ratios were recorded at 30 d and at 60 d.
The bulbing ratio was calculated by dividing the
maximum plant leaf base diameter by the minimum neck
diameter. Plant leaf base and neck diameters were
measured with vernier callipers. A bulbing ratio equal to
or greater than 2.0 was taken as indicative of definite
bulbing (Terabun, 1965; Kedar et al., 1975; Wright and
Sobeih, 1986; Brewster et al., 1987).
Random samples of two plants from the extra plants in
each treatment were dissected to observe inflorescence
initials using a binocular microscope. The samples were
fixed in 70% (v/v) methanol and kept in a refrigerator
prior to processing for scanning electronic microscopy
(SEM). The time to bulb maturation was recorded when
neckfall started. The number of leaves was recorded at
the time of inflorescence initiation.
Analysis of variance was performed using SAS
software (SAS, 1985). Regression analysis was
performed with Microsoft EXCEL following the
procedure of Gomez and Gomez (1984). Fitted planes
from multiple regression analyses were plotted by 3-D
graphs using ‘SlideWrite’ Version 2.0 (1989).
Inflorescence initiation in ‘Phulkara’ took place only at a
photoperiod of 8 h d–1. Therefore, multiple regression
analysis for the time taken to inflorescence initiation was
not possible, and 3-D graphs could not be plotted for this
cultivar. However, the results are described and
discussed.
RESULTS
Bulbing ratio
Bulbing ratios increased curvi-linearly with increasing
temperature and lengthening photoperiod for all
cultivars examined (Figure 1A–P). For example, after
60 d, the bulbing ratio under a 14 h d–1 photoperiod
Country of origin Day-length status Season
Netherlands Long Spring
Netherlands Intermediate Spring
Japan Long Autumn
Turkey Intermediate Spring
UK Long Spring
Australia Intermediate Spring
Netherlands Long Spring
Pakistan Short Autumn
490 Bulbing and bolting in onion cultivars

FIG. 1
Effect of temperature, photoperiod and time on the bulbing ratio in eight onion cultivars of diverse origin at 30 d and 60 d (Panels A – P).
2
(r = 0.72 – 0.91).
K. M. KHOKHAR 491
492 Bulbing and bolting in onion cultivars
increased from 2.07 and 1.50 at 13.7°C to 2.95 and 2.51 at
29.2°C for ‘Senshyu Yellow’ and ‘Jaune Espagnol’,
respectively. Similarly, after 60 d, the bulbing ratio at
29.2°C increased from 1.42 and 1.33 with an 8 h d–1
photoperiod, to 4.20 and 3.42 with a 17 h d–1 photoperiod
for ‘Senshyu Yellow’ and ‘Jaune Espagnol’, respectively.
For ‘Senshyu Yellow’, ‘Jaune Espagnol’, ‘Australian
Brown’ and ‘Phulkara’ there was a significant interaction
(P < 0.001) between temperature and photoperiod, such
that the bulbing ratio was maximised under a
photoperiod of 17 h d–1 at a mean temperature of 25°C.
In ‘Hygro’ and ‘Delta’, there was also a significant
interaction (P < 0.001) between photoperiod and time,
such that the effect of photoperiod was more
pronounced after 60 d than after 30 d. In ‘Sito’, there was
a significant interaction (P < 0.001) between
temperature, photoperiod and time, such that the
effect of temperature and photoperiod on the bulbing
ratio was more pronounced after 60 d than after 30 d.
There was an interaction between temperature and time
in ‘Rijnsburger Balstora’, such that the effect of
temperature on the bulbing ratio was more pronounced
after 60 d than after 30 d.
‘Senshyu Yellow’, ‘Jaune Espagnol’, ‘Hygro’, ‘Sito’,
‘Delta’, ‘Australian Brown’ and ‘Rijnsburger Balstora’
(intermediate to LD types), bulbed under 14 h d–1 and
17 h d–1 photoperiods. The short-day (SD) cultivar,
‘Phulkara’ (Pakistan), bulbed under photoperiods of
11 h d–1, 14 h d–1 and 17 h d–1. However, no bulbing was
observed under an 8 h d–1 photoperiod in any cultivar, at
any mean temperature. In all cultivars except ‘Sito’,
bulbing occurred earlier (i.e., 30 d) after exposure to the
different inductive photo-thermal regimes. For example
in ‘Senshyu Yellow’ and ‘Phulkara’, bulbing occurred
after 30 d at 11°C under a 17 h d–1 photoperiod, and from
14° – 29°C under 14 h d–1 and 17 h d–1 photoperiods,
while ‘Phulkara’ also bulbed under an 11 h d–1
photoperiod from 25° – 29°C. Bulbing in ‘Jaune
Espagnol’, ‘Hygro’ and ‘Rijnsburger Balstora’ occurred
after 30 d at temperatures from 25° – 29°C under a 17 h
d–1 photoperiod, but ‘Hygro’ also bulbed from 19° – 23°C
at a 17 h d–1 photoperiod. ‘Delta’ bulbed from 23° – 29°C
under 14 h d–1 and 17 h d–1 photoperiods, and also at 19°C
under a 17 h d–1 photoperiod. In ‘Australian Brown’,
bulbing also occurred after 30 d from 19° – 29°C under
14 h d–1 and 17 h d–1 photoperiods, and from 11° – 14°C
under a 17 h d–1 photoperiod. However ‘Sito’ was late,
and bulbing was observed only after 60 d at 25° – 29°C
under 14 h d–1 and 17 h d–1, and at 20° – 23°C under 17 h
d–1 photoperiods. Bulbing occurred faster under a longer
photoperiod (17 h d–1) and at higher temperatures in all
cultivars. In ‘Jaune Espagnol’, bulbing occurred after 30 d
at 17 h d–1 at higher temperatures from 25º – 29ºC, but
after 60 d under 14 h d–1, and 17 h d–1 from 13º – 29ºC.
Time to floral initiation
Under low temperatures, all genotypes tended to
initiate flowers rather than to bulb. Time to floral
initiation in ‘Senshyu Yellow’, ‘Hygro’ and ‘Delta’
decreased linearly with decreasing temperature and
increasing photoperiod (Figure 2A, C, E). In ‘Jaune
Espagnol’, ‘Sito’, ‘Australian Brown’ and ‘Rijnsburger
Balstora’, time to floral initiation also decreased linearly
with decreasing temperature, but decreased curvi-
linearly with increasing photoperiod (Figure 2B, D, F, G).
Thus, time to floral initiation under an 8 h d–1
photoperiod decreased from 185 d and 175 d at 13.1°C,
to 183 d and 170 d at 10.9°C for ‘Senshyu Yellow’ and
‘Jaune Espagnol’, respectively. Time to floral initiation at
13.1°C decreased from 185 d and 175 d with an 8 h d–1
photoperiod, to 181 d and 168 d with an 11 h d–1
photoperiod for ‘Senshyu Yellow’ and ‘Jaune Espagnol’,
respectively. Adding an interaction term did not
significantly improve these relationships.
‘Senshyu Yellow’, ‘Jaune Espagnol’, ‘Hygro’, ‘Sito’,
‘Delta’, ‘Australian Brown’ and ‘Rijnsburger Balstora’
(intermediate to LD types), initiated flowers under 8 h
d–1 and 11 h d–1 at temperatures from 11° – 13°C. In
‘Jaune Espagnol’, ‘Hygro’, ‘Sito’, ‘Australian Brown’ and
‘Rijnsburger Balstora’, inflorescences were also initiated
under 14 h d–1 at mean temperatures from 11° – 13°C.
‘Rijnsburger Balstora’ initiated inflorescences even at
relatively warm temperatures (19°C) under
photoperiods of 11 h d–1 and 14 h d–1. However, in the
short-day ‘Phulkara’, inflorescence initiation occurred
only at a photoperiod of 8 h d–1 at temperatures from
11° – 13°C.
Time to bulb maturity
The time to bulb maturity decreased linearly with
increasing temperature and also with lengthening
photoperiod in all cultivars (Figure 3A–H). For example,
for ‘Senshyu Yellow’ and ‘Australian Brown’, time to
bulb maturity at a 14 h d–1 photoperiod decreased from
98 d and 90 d at 22.7°C, and to 67 d and 61 d at 29.1°C,
respectively. Similarly, time to bulb maturity at 29.1°C
decreased from 67 d and 60 d at a 14 h d–1 photoperiod,
and to 51 d and 55 d at a 17 h d–1 photoperiod for
‘Senshyu Yellow’ and ‘Rijnsburger Balstora’,
respectively. However, there was an interaction between
photoperiod and temperature such that increasing
photoperiods decreased the time to bulb maturity under
all temperatures, but the effect of photoperiod was more
pronounced at lower temperatures. The short-day
‘Phulkara’ was earliest to mature, taking 49 d to mature
under a 17 h d–1 photoperiod at a temperature of 29°C,
whereas other cultivars (intermediate to LD type)
matured in 51 – 55 d under the same photo-thermal
regime.
DISCUSSION
The results of these experiments show that the
dynamics of bulbing and bolting in onion plants are
influenced significantly by photoperiod and
temperature. Increasing photoperiods promoted bulbing,
but, under short photoperiods (8 h d–1), plants did not
bulb even after a long period of growth. This confirms
previous findings (Heath, 1943; Heath and Mathur, 1944;
Austin, 1972; Kedar et al., 1975; Steer, 1980) that
increasing photoperiod promotes bulbing, and short
photoperiods delay or, if sufficiently short, completely
inhibit bulbing. Increasing temperatures promoted
bulbing and hastened maturity. This strengthens previous
findings (Heath, 1945; Kato, 1964) that LD and higher
temperatures cause earlier bulbing and maturity. It was
shown earlier that bulbing did not occur at lower
temperatures of 10° – 15.5°C (Thompson and Smith,
 K. M. KHOKHAR 493

FIG. 2
Effect of temperature and photoperiod on the time to floral initiation in seven onion cultivars of diverse origin (Panels A – G). r2 = 0.96 – 0.99.

= no response,  = response.
494 Bulbing and bolting in onion cultivars

FIG. 3
Effect of temperature and photoperiod on the time to neckfall in eight onion cultivars of diverse origin (Panel A – H). r2 = 0.90 – 0.98,
= no
response,  = response.
 K. M. KHOKHAR
1938); however, the results of our experiments show that,
in bolting-resistant cultivars, bulbing can take place at
lower temperatures (11° – 13.8°C) under a longer
photoperiod (17 h d–1). However, the present studies
support the findings of Heath (1943; 1945), who reported
that lower temperatures could greatly delay or prevent
bulbing. With an increase in temperature, bulb
development under long photoperiods was accelerated;
while, under short photoperiods (8 – 11 h d–1), no bulbing
occurred even at warm temperatures. This is in
agreement with previous reports of other workers
(Heath, 1943; 1945; Kato, 1964; Butt, 1968; Steer, 1980;
Terabun, 1980) who showed that, with an increase in
temperature, the rate of bulb development increases;
but, if day-lengths are sufficiently short, no bulbing will
occur, even at warm temperatures.
Under low temperatures, the time to floral initiation
was hastened with decreasing temperatures and
lengthening photoperiods (8 – 14 h d–1). This confirms
previous findings (Shishido and Saito, 1975; Brewster,
1983) that extended photoperiods under low inducing
temperatures promoted inflorescence initiation. The
present studies confirm the findings of Holdsworth and
Heath (1950), van Kampen (1970), Brewster (1982b) and
Bertaud (1988) that inflorescence development after
initiation is positively correlated with photoperiod, but
are in conflict with some reports (Holdsworth and
Heath, 1950; van Kampen, 1970) indicating no effect of
photoperiod on inflorescence initiation.
Our studies have shown that inflorescences in most
onion cultivars (‘Senshyu Yellow’ ‘Delta’ ‘Jaune
Espagnol’, ‘Hygro’, ‘Sito’, ‘Australian Brown’) can
initiate under short-to-intermediate day-lengths (8 h d–1,
11 h d–1 or 14 h d–1), at temperatures from 11° – 13°C.
Initiation of inflorescences in ‘Rijnsburger Balstora’ can
take place under a short photoperiod (8 h d–1) at
temperatures from 11° – 13°C, and under short-to-
intermediate day-lengths (11 h d–1 and 14 h d–1) from
11° – 19°C. ‘Phulkara’ can initiate inflorescences only
under short-days (8 h d–1) at temperatures from
11° – 13°C. Inflorescences in most cultivars initiated at
11° – 13°C after 154 – 184 d, except in ‘Rijnsburger
Balstora’, where initiation took place at 11° – 18.9°C
after 162 – 185 d. This confirms the findings of Shishido
and Saito (1975, 1977), Brewster (1983), Bertaud (1988),
De Bon and Rhino (1988) and Peters (1990) that low
temperatures between 7° – 13°C for 20 d – 90 d were
optimum for flower induction in most onion cultivars
and that bolting-resistant varieties required a longer cold
stimulus compared to normal Spring-sown cultivars.
Most cultivars in the present study were bolting-resistant
and required a longer cold stimulus (154 – 185 d) for
flower induction. ‘Rijnsburger Balstora’ also initiated
495
inflorescences at relatively higher temperatures (18.9°C)
than the other cultivars studied here. Cultivars adapted
to warmer conditions have a lesser requirement for cold,
and possibly a higher optimum vernalisation
temperature. This could be attributed to genotype, and
confirms reports (Sinnadurai, 1970) that certain cultivars
(for example ‘Bawku’) flowered at ambient night
temperature of 15° – 21°C.
These studies have clearly shown that photoperiod
and temperature are extremely important environmental
variables for flowering and bulbing in onion, and that
different cultivars show variations in their responses.
Considerable variability exists in the global
environment. It is therefore possible to identify suitable
photo-thermal regimes for flowering or bulbing for a
particular cultivar in a particular region. It is also
possible to combine the fitted relationships for bulbing
and flowering for each cultivar to have the full photo-
thermal response for flowering and bulbing.
Flowering in ‘Senshyu Yellow’ and ‘Delta’ will occur
under shorter photoperiods (8 h d–1 and 11 h d–1) at
temperatures from 11° – 13°C, while bulbing will occur
under intermediate-to-LD (14 h d–1 and 17 h d–1) at
temperatures from 11° – 29°C. ‘Jaune Espagnol’, ‘Hygro’
and ‘Sito’ will initiate flowers under short-to-
intermediate day-lengths (8 h d–1, 11 h d–1 and 14 h d–1)
at mean temperatures from 11° – 13°C. However, these
will bulb under intermediate day-length (14 h d–1) at
temperatures from 20° – 29°C. Under a long
photoperiod (17 h d–1), bulbing will occur at
temperatures from 11° – 29°C. ‘Australian Brown’ will
flower under short photoperiods (8 h d–1 and 11 h d–1) at
temperatures from 11° – 13°C, and under intermediate
photoperiods (14 h d–1) at 11°C. However, the latter will
bulb under intermediate day-lengths (14 h d–1) at
temperatures from 14° – 29°C. Under a long
photoperiod (17 h d–1), bulbing will occur from
11° – 29°C. Flowering in ‘Rijnsburger Balstora’ will
occur under short photoperiod (8 h d–1) at temperatures
from 11° – 13 °C and under short-to-intermediate day-
lengths (11 h d–1 and 14 h d–1) from 11° – 19°C. However,
it will bulb under intermediate day-lengths (14 h d–1) at
temperatures from 23° – 29°C, and under a long
photoperiod (17 h d–1) from 11° – 29°C. ‘Phulkara’ will
flower under short-day (8 h d–1) at temperatures from
11° – 13°C. However, it will bulb under a wide range of
photoperiods (11 h d–1, 14 h d–1 and 17 h d–1) at
temperatures from 11° – 29°C. The bulbing ratio, bulb
maturity, and floral initiation of onion plants are largely
dependent on temperature and photoperiod. Thus, plant
development can be modified towards bulbing, or
towards flowering under suitable photo-thermal
regimes.
 496 Bulbing and bolting in onion cultivars
REFERENCES
AUSTIN, R. B. (1972). Bulb formation in onions as affected by pho-
toperiod and spectral quality of light. Journal of Horticultural
Science, 47, 493–504.
BERTAUD, D. S. (1988). Effects of chilling duration, photoperiod and
temperature on floral initiation and development in sprouted
and unsprouted onion bulbs. In: Proceedings of the 4th
EUCARPIA Allium Symposium, Institute of Horticultural
Research, Wellesbourne, UK. 245–261.
BREWSTER, J. L. (1982a). Flowering and seed production in over-
wintered cultivars of bulb onions. I. Effects of different raising
environments, temperatures and daylengths. Journal of
Horticultural Science, 57, 93–101.
BREWSTER, J. L. (1982b). Flowering and seed production in over-
wintered cultivars of bulb onions. II. Quantitative relationships
between mean temperatures and daylengths and the rate of
inflorescence development. Journal of Horticultural Science, 57,
103–108.
BREWSTER, J. L. (1983). Effects of photoperiod, nitrogen nutrition
and temperature on inflorescence initiation and development
in onion (Allium cepa L.). Annals of Botany, 51, 429–440.
BREWSTER, J. L. (1987a). The effect of temperature on the rate of
sprout growth and development within stored onion bulbs.
Annals of Applied Biology, 111, 463–467.
BREWSTER, J. L. (1987b). Vernalization in the onion - a quantitative
approach. In: Manipulation of Flowering. (Atherton, J.G., Ed.).
Butterworths, London. UK. 171–183.
BREWSTER, J. L., LAWES, W. and WHITLOCK, A. J. (1987). The phe-
nology of onion bulb development at different sites and its rel-
evance to incomplete bulbing (‘thick-necking’). Journal of
Horticultural Science, 62, 371–378.
BUTT, A. M. (1968). Vegetative growth, morphogenesis and carbo-
hydrate content of the onion plant as a function of light and
temperature under field and controlled conditions.
Mededelingen Landbouwhogeschool Wageningen, 68, 94–191.
DAVIS, G. N. and JONES, H. A. (1944). Experiments with the trans-
plant onion crop in California. California Agricultural
Experiment Station Bulletin No. 682, Davis, CA, USA. 20 pp.
DE BON, H. and RHINO, B. (1988). Flowering in the various cultivars
of onions (Allium cepa L.) brought to Martinique (French West
Indies). In: Proceedings of the 4th EUCARPIA Allium
Symposium, Institute of Horticultural Research, Wellesbourne,
UK. 262–266.
DOWKER, B. D. (1990). Onion breeding. In: Onions and Allied
Crops. Volume I. (Rabinowitch, H.D. and Brewster, J.L., Eds.).
CRC Press Inc., Boca Raton, FL, USA. 215–232.
GOMEZ, K. A. and GOMEZ, A. A. (1984). Statistical Procedures for
Agricultural Research. 2nd Edition. John Wiley and Sons, Ltd.,
New York, USA. 680 pp.
HARTSEMA, A. M. (1947). The periodical development of the onion
(Allium cepa L.), var. Giant Zittau. Mededelingen
Landbouwhogeschool Wageningen, 48, 265–300. (In Dutch with
English summary).
HEATH, O. V. S. (1943). Studies on the physiology of the onion plant.
I. An investigation of factors concerned in the flowering
(bolting) of onions grown from sets and its prevention. Part 2.
Effects of day length and temperature on onions grown from
sets and general discussion. Annals of Applied Biology, 30,
308–322.
HEATH, O. V. S. (1945). Formative effects of environmental factors
as exemplified in the development of the onion plant. Nature,
155, 623–626.
HEATH, O. V. S. and MATHUR, P. B. (1944). Studies on the physiology
of the onion plant. II. Inflorescence initiation and development,
and other changes in the internal morphology of onion sets, as
influenced by temperature and day length. Annals of Applied
Biology, 31, 173–187.
HEATH, O. V. S. and HOLDSWORTH, M. (1948). Morphogenic factors
as exemplified by the onion plant. In: Growth. (Daneellie, J.F.
and Brown, R., Eds.). Symposia of the Society for Experimental
Biology, 2, 326–350.
HOLDSWORTH, M. and HEATH, O. V. S. (1950). Studies on the physi-
ology of the onion plant. Journal of Experimental Botany, 1,
353–375.
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ITO, K. (1956). Studies on the bolting of onion. I. Relation between
the flower bud formation and the bulb division. Journal of the
Japanese Society for Horticultural Science, 25, 187–193. (In
Japanese with English summary).
KATO, T. (1964). Physiological studies on bulb formation and dor-
mancy in onion plant. III. Effects of external factors on the bulb
formation and development. Journal of the Japanese Society for
Horticultural Science, 33, 53–61. (In Japanese with English
summary).
KEDAR, N., LEVY, D. and GOLDSCHMIDT, E. E. (1975). Photoperiodic
regulation of bulbing and maturation of ‘Bet Alpha’ onions
(Allium cepa L.) under decreasing daylength conditions.
Journal of Horticultural Science, 50, 373–380.
PETERS, R. (1990). Seed production in onions and some other
Allium species. In: Onions and Allied Crops. Volume I.
(Rabinowitch, H. D. and Brewster, J. L., Eds.). CRC Press Inc.,
Boca Raton, FL, USA. 161–176.
PIKE, L. M. (1986). Onion breeding. In: Breeding Vegetable Crops.
(Bassett, M.J., Ed.). AVI Publishing Co., Westport, CT, USA.
357–394.
RABINOWITCH, H. D. (1985). Onions and other edible Alliums. In:
CRC Handbook of Flowering. Volume I. (Halevy, A.H., Ed.).
CRC Press Inc., Boca Raton, FL, USA. 398–405.
RABINOWITCH, H. D. (1990). Physiology of flowering. In: Onions and
Allied Crops. Volume I. (Rabinowitch, H.D. and Brewster, J.L.,
Eds.). CRC Press Inc., Boca Raton, FL, USA. 133–134.
S.A.S. (1985). Statistical Analysis System User’s Guide: Statistics. SAS.
Inc., Cary, NC, USA. (http://www.asu.edu/sas/sasdoc/sashtml/).
SHISHIDO, Y. and SAITO, T. (1975). Studies on the flower bud forma-
tion in onion plants. I. Effects of temperature, photoperiod and
light intensity on the low temperature induction of flower buds.
Journal of the Japanese Society for Horticultural Science, 44,
122–130. (In Japanese with English summary).
SHISHIDO, Y. and SAITO, T. (1976). Studies on the flower bud forma-
tion in onion plants. II. Effects of physiological conditions on
the low temperature induction of flower buds on green plants.
Journal of the Japanese Society for Horticultural Science, 45,
160–167. (In Japanese with English summary).
SHISHIDO, Y. and SAITO, T. (1977). Studies on the flower bud forma-
tion in onion plants. III. Effects of physiological conditions on
the low temperature induction of flower buds in bulbs. Journal
of the Japanese Society for Horticultural Science, 46, 310–316.
(In Japanese with English summary).
SINNADURAI, S. (1970). A note on the bulbing and flowering habit of
the Bawku onion. Tropical Agriculture (Trinidad), 47, 77–79.
SLIDE WRITE PLUS (1989). Advanced Graphics Software Inc.,
Sunnyvale, CA, USA. (http://www.slidewrite.com/TrialGuide.asp).
STEER, B. T. (1980). The bulbing response to daylength and temper-
ature of some Australian cultivars of onion (Allium cepa L.).
Australian Journal of Agricultural Research, 31, 511–518.
TERABUN, M. (1965). Studies on the bulb formation in onion plants.
I. Effects of light quality on the bulb formation and the growth.
Journal of the Japanese Society for Horticultural Science, 34,
196–204. (In Japanese with English summary).
TERABUN, M. (1980). Effect of cyclic lighting on photoperiodic
responses in onion plant. Journal of the Horticultural
Association of Japan, 49, 375–379.
THOMPSON, H. C. and SMITH, O. (1938). Seedstalk and bulb devel-
opment in the onion (Allium cepa L.). Cornell University
Agricultural Experiment Station Bulletin, No. 708. 21 pp.
VAN KAMPEN, J. (1970). Shortening the breeding cycle in onions.
Mededelingen Proefstation Voor de Groenteteelt in de
Vollegrond in Nederland. 51, 72. (In Dutch with English
summary).
WRIGHT, C. J. and SOBEIH, W. Y. (1986). The photoperiodic regula-
tion of bulbing in onions (Allium cepa L.). I. Effects of irradi-
ance. Journal of Horticultural Science, 61, 331–335.
WOODBURY, G. W. (1950). A study of factors influencing floral initi-
ation and seedstalk development in the onion, Allium cepa
Linn. Idaho Agricultural Experiment Station Research Bulletin,
No. 18. 27 pp.