Annals of Botany 78 : 423430, 1996

Bulbing in Onions : Photoperiod and Temperature Requirements and Prediction of

Bulb Size and Maturity
J. E. L A N C A S T ER*, C. M. T R I G G S, J. M. D E R U I T E R* and P. W. G A N D AR
* New Zealand Institute Crop & Food Research, Priate Bag 4704, Christchurch
Statistics Dept, Uniersity of Auckland, Priate Bag 92019, Auckland, and
New Zealand Institute for Horticulture and Food Research, Priate Bag, Palmerston North, New Zealand

Received : 24 April 1995 Accepted : 27 February 1996

Bulb size and maturity are key characteristics of an onion crop and the onset of bulbing is an important determinant
of these. In this paper we describe an experiment in which bulb and neck diameter and leaf number were measured
in onion crops (cultivars Pukekohe Longkeeper and Early Longkeeper) with different sowing dates planted at two
different locations in New Zealand. A sensitive indicator of earliest time of bulbing was developed using the ratio of
bulb and neck diameters and the statistical technique of cusums. Bulb diameter at bulbing was related to thermal time
accumulated prior to bulbing. Bulbing only occurred when dual thresholds of a minimum thermal time of 600 degree
days and a photoperiod of 1375 h were reached. Mathematical relationships were developed between leaf number,
sowing date, bulbing date and bulb growth and maturity. Final bulb size could be predicted from bulb size at bulbing
and number of leaves produced after bulbing. Bulb maturity date could be predicted by number of leaves after
bulbing. # 1996 Annals of Botany Company

Key words : Allium cepa L., onion, temperature, photoperiod, bulb-neck ratio, leaf number, bulbing.

density (Rogers, 1977). In the field experiments described

INTRODUCTION
The onion (Allium cepa L.) is a biennial plant, and the bulb
is a vegetative overwintering stage in the life cycle of the
plant. An onion plant is composed of leaves which arise
alternately from a small flattened stem, or base plate, so that
older leaves are on the outside and younger leaves on the
inside of the stem. Each leaf is composed of a photosynthetic
leaf blade and a non-photosynthetic, storage leaf base
(scale). During the growth of the plant the leaf scales
thicken and form the characteristic bulb. Onions have a
determinate growth habit. At the onset of bulbing, leaf
sheaths swell, bladeless bulb scales are initiated and these
swell to form the central storage tissue of the bulb. Leaf
blades initiated prior to bulbing develop to full expansion
and the green top lodges at top down . Mature onion bulbs
can range in size from 5 mm to over 100 mm bulb diameter.
Bulb formation and subsequent growth are influenced by
temperature and photoperiod (Brewster, 1977). Bulb form-
ation is promoted by long days and high temperatures
(Magruder and Allard, 1937 ; Heath, 1945 ; Kato, 1964).
Steer (1980 b) showed the importance of night temperature
in the induction of bulbing in phytotron-grown plants and
Lercari (1984) and Austin (1972) showed that far-red light
was also necessary. In New Zealand onions are sown at low
densities so that light quality is not a limiting factor.
Clearly, many factors other than temperature and photo-
period affect bulb growth), e.g. irrigation (de Lis et al., 1967 ;
van Eeden and Myburgh, 1971 ; Chaudhry and Erinne,
1984), fertiliser application (Hassan and Ayoub, 1978),
weed competition (Hewson and Roberts, 1973) and planting
0305-7364}96}10042308 $18.00}0
below weed competition, density, fertilizer and water were
non-limiting.
Although temperature and photoperiod are known to
interact to induce bulbing (Brewster, 1990) there has been
little work in which the photothermal requirements for
bulbing in the field have been specified. De Ruiter (1986)
examined the relationship between bulb size and thermal
time from emergence to harvest for six cultivars at different
sowing dates. The relative growth rates of bulbs were
linearly related to accumulated thermal time. However, the
responses for individual cultivars were different. Prediction
of maturation time and the effects of photoperiod were not
defined precisely in this study.
Prediction of the onset of bulb development is required
for better prediction of maturity dates. Predictive models
for estimating the rate of bulb development and final bulb
size would also be of value. In this paper we describe an
experiment in which bulb and neck diameter and leaf
number were measured in onion crops with different sowing
dates planted at two locations. We show that bulbing is
determined by the dual thresholds of temperature and
photoperiod and we discuss the implications of these results
for predicting bulb size and maturity.
MATERIALS AND METHODS
Plant material
The cultivars Pukekohe Longkeeper and Early Longkeeper
were grown at Pukekohe (3712 S) in 1986}87 and 1987}88
# 1996 Annals of Botany Company
424 Lancaster et al.Photoperiod and Temperature Effects on Bulbing in Onions
T 1. Bulb characteristics of Pukekohe Longkeeper (PLK) and Early Longkeeper (ELK) onions planted on sowing dates
between May and Sep. in Pukekohe, 198687

Median bulbing Median diameter Thermal time emergence to Mean bulb diameter at
Site Planting date date of bulbing (mm) bulbing (degree days) harvest (mm)

PLK 1 9 Jun. 9 Nov. 132 818 677

2 16 Jun. 28 Oct. 82 696 606
3 17 Jun. 9 Nov. 101 789 755
4 3 Jul. 6 Nov. 86 709 636
5 4 Jul. 8 Nov. 56 729 633(*)
6 4 Jul. 1 Nov. 101 662 691(*)
7 17 Jul. 6 Nov. 89 646 648
8 18 Jul. 31 Oct. 8 579 597
9 20 Jul. 6 Nov. 67 627 537(*)
10 24 Jul. 6 Nov. 59 609 567
11 19 Aug. 16 Nov. 59 663 538
12 19 Aug. 24 Nov. 52 582 584
13 24 Sep. (19 Dec.) (! 20) 664 395
ELK 14 30 May 1 Nov. 111 792 602
15 30 May 28 Oct. 118 760 *
16 9 Jun. 31 Oct. 128 733 648
17 9 Jun. 2 Nov. 131 751 585
18 12 Jun. 30 Oct. 109 711 677
19 14 Jun. 29 Oct. 11 696 565
20 15 Jun. 4 Nov. 98 743 515
21 17 Jun. 29 Oct. 82 688 598
22 19 Jun. 31 Oct. 89 695 706
23 3 Jul. 26 Oct. 92 600 561
24 17 Jul. 6 Nov. 9 637 509
25 19 Aug. 20 Nov. 45 612 551
26 24 Sep. (19 Dec.) (! 20) 664 364

Note : All sampled onions from site 15 lost. (*) Many sampled onions from site 5, 6 and 9 lost.

T 2. Bulb characteristics of Pukekohe Longkeeper (PLK) and Early Longkeeper (ELK) onions planted on sowing dates
between May and Sep. in Pukekohe, 198788

Median bulbing Median diameter Thermal time emergence to Mean bulb diameter at
Site Planting date date of bulbing (mm) bulbing (degree days) harvest (mm)

PLK 27 4 May 9 Nov. 116 1014 606

28 7 Jun. 2 Nov. 16 819 748
29 9 Jul. 3 Nov. 132 659 677
30 9 Jul. 1 Nov. 114 639 62
31 10 Jul. 4 Nov. 94 662 776
32 21 Jul. 23 Nov. 146 792 577
33 31 Jul. 21 Nov. 103 729 624
34 12 Aug. 30 Nov. 103 733 72
35 15 Aug. 28 Nov. 72 657 452
36 29 Aug. 1 Dec. 103 626 571
37 25 Sep. 18 Dec. 103 600 552
ELK 38 4 May 30 Oct. 101 1033 584
39 26 May 25 Oct. 125 835 635
40 28 May 25 Oct. 127 822 649
41 28 May 29 Oct. 146 866 667
42 28 May 26 Oct. 136 834 683
43 7 Jun. 1 Nov. 133 844 703
44 12 Jun. 6 Nov. 12 851 621
45 23 Jun. 27 Oct. 137 701 738
46 7 Jul. 4 Nov. 128 706 72
47 11 Jul. 30 Oct. 106 647 813
48 12 Aug. 27 Nov. 91 744 646
49 25 Sep. 14 Dec. 78 574 517

and at Palmerston North (4012 S) in 1985}86 (Tables Sep. in 1987}88 and from the 31 May to 25 Sep. in 1986}87.
13). These sites were commercial production fields except for
In Pukekohe, sowing dates ranged from the 4 May to 25 sites (Table 1 ; 1, 7, 12, 13, 16, 2426) (1986}87), (Table 2 ; 27,
 Lancaster et al.Photoperiod and Temperature Effects on Bulbing in Onions
T 3. State of the plant with respect to bulbing
Thermal time
Photoperiod (h) ! 600 & 600
! 1375 OFF OFF
& 1375 OFF ON
34, 37, 38, 48, 49) (1987}88) which were at the Pukekohe
research station and were managed as part of an onion
breeding programme. The latter 14 sites were chosen to
extend the range of planting dates. In Pukekohe seeds where
sown directly into the field and grown according to standard
commercial practices (Lancaster et al., 1995). Data from an
earlier trial at Palmerston North are also used in this paper
(de Ruiter, 1986). In this trial sowing dates from 14 May to
28 Aug. 1986 were used. The trial was a split plot randomized
complete block design with three replications. Seeds were
sown directly and thinned to 10 cm spacing within rows. At
sowing, 40 t ha" of 30 % potassic superphosphate was
applied along with 108 kg ha" of urea. Two subsequent
applications of urea (2108 kg ha") were applied at 8 week
intervals.
Measurement of onions
At Pukekohe, 30 onions were randomly selected at each
site and labelled individually. The diameter of the neck and
bulb of each onion was measured with vernier callipers
fortnightly from the beginning of Sep. until the end of Oct.
425
diameter of the bulb begins to increase very rapidly and so
the ratio of diameters also increases. A sensitive, reliable
and non-destructive indicator is required in order to detect
the commencement of bulbing on individual plants. The
technique of taking cumulative sums (cusums) can be used
to detect such change-points in a series of measurements
over time (British Standard 5703 ; Woodward and Gold-
smith, 1964 ; Lancaster, Triggs and Barrett, 1986). We used
the cusum method, with a threshold ratio of 12, to detect
the onset of bulbing. This threshold ratio was chosen to take
account of small variations in the measurement of the
diameters, particularly when both measurements are less
than 5 mm. This threshold value was subtracted from the
calculated ratio for each bulb at each time of measurement,
and the differences accumulated. The cumulative sums, or
cusums, of these differences were then plotted against time
(Fig. 1).
Before bulbing the ratio fluctuates about a value of one
(Fig. 1 A), and the cusum plot is close to a straight line (Fig.
1 B). After bulbing the ratio increases rapidly and so the
cusum becomes positive and increases very rapidly. The fact
that the differences from the target ratio are accumulated,
means that the cusum plot is very sensitive to the changes in
ratio. Dates of bulbing can be estimated using the minimum
points when the cusum plot increases sharply.
Examination of site records suggested that 50 % seedling
emergence took place approximately 18 d after planting for
most sites. Thermal time (growing degree days above 5 C
air temperature) was accumulated between the date of
emergence and date of bulbing for each site. Thermal time,
D, was calculated as :

and weekly thereafter until harvested. Diameters were only
measured when they were greater than 2 mm.
At Palmerston North, plants were monitored for leaf
numbers appearance and bulb diameter. The leaf tips of five
plants per plot were marked with paint at 2 week intervals
throughout growth, beginning at the appearance of the
second true leaf. The total number of emerged leaves and
the total numbers of leaves per plant were recorded at
similar intervals. The number of leaves present at any time
is determined by the balance between the rate of appearance
of new leaves and the rate of loss of old leaves. At top down,
plants were harvested and measurements made of total
number of emerged leaves, total leaves and scales initiated.
Climate measurements
At Pukekohe, climatic data (rainfall, maximum and
minimum air temperature) was recorded at the meteoro-
logical site located within 20 km of all of the trial sites. At
Palmerston North, daily maximum, minimum and hourly
mean temperatures for air and soil (2, 5 and 10 cm depth)
were recorded for the duration of the experiment. Day
lengths were calculated using the method of Keisling (1982).
Data analysis
During the early growth of the onion plant the ratio of
bulb to neck diameter is close to one. At bulbing the
9 :
n (T maxT min) +
D 3 i i Tb
i="
2
where T max and T min are daily maximum and minimum
temperature, T b is the base temperature and n is number of
days between emergence and bulbing. The plus sign indicates
that the summation only included days when mean
temperature exceeded the base temperature. A minimum
variance method (Arnold, 1959), was used to select the
appropriate base temperature for rate of appearance of
leaves 39. A linear rate of leaf appearance was assumed in
calculating the base temperature. The estimated base
temperature for the two cultivars was very close to 5 C, so
this value was used in subsequent calculations.
Statistical analysis
Although repeated measurements were made on in-
dividual bulbs, the analyses and models in this paper pertain
to the behaviour of average plants in each site. Thus, mean
and median values for bulb size and leaf number have been
used in all analyses. Regression models were fitted using the
MINITAB statistical package. A residual mean square
error statistic was calculated for model predictions and
observed values for bulb size and maturity date using
RMSD '3 (Observedpredicted)
n
#
426 Lancaster et al.Photoperiod and Temperature Effects on Bulbing in Onions
9 1100
A

8 1000
Ratio bulb diameter : neck diameter

7 900

Thermal time (D)

6 800

5 700

600
4

500
3 12 13 14 15
Photoperiod (h)
400
2 260 280 300 320 340 360
Day of bulbing
1 Sep. Oct. Nov. Dec.

F. 2. Thermal time in degree days and calender date of bulbing for

Pukekohe Longkeeper (E) and Early Longkeeper (*) onions grown at
various sowing dates in Pukekohe in 198687 and 198788. The dotted
B lines indicate the derived critical minima for thermal time and
photoperiod.
20

sites ranged from 28 Oct. to 24 Nov. in 1986}87 and from

1 Nov. to 18 Dec. with a median over the two seasons of 9
15 Nov. Bulbing had commenced prior to the first measurement
for both ELK and PLK at the extremely late planted sites,
13 and 26, in 1986}87. In these cases, bulb diameters were
Cusums (1.2)

10 less than 2 mm so that no reliable data for date of bulb

5
0 early planted sites (Tables 1 and 2). Excluding the very late
5
Sep. Oct.
Date
F. 1. Estimation of date of bulbing. A, Examples of the change in
bulb neck ratio with time of five onions. B, Corresponding changes in
cusums of this ratio.
Goodness of fit was evaluated using r# and a standardized
value of RMSD relative to the mean observed value.
RESULTS
Date of bulbing and plant size
Median bulbing dates of the 30 plants at each Pukekohe
site, calculated using the cusum method (Fig. 1), are given
in Tables 1 and 2. Estimated bulbing dates for ELK sites
ranged from 26 Oct. to 20 Nov. in 1986}87 and from 25 Oct.
to 14 Dec. in 1987}88 with a median over all plantings in the
two seasons of 30 Oct. The corresponding dates for the PLK
initiation were available. The dates of 30 Oct. and 9 Nov.
correspond respectively to day lengths of 135 h and 14 h
(see below). This suggests that daylengths of at least 135 h
are implicated in bulbing.
Bulb diameter for each site at the median bulbing date
ranged from less than 2 mm to 160 mm between late and
planted sites (13, 26) the smallest median diameter at
bulbing was 45 mm.
Nov. Dec.
Physiological time to bulbing
At the Pukekohe sites in both seasons, thermal time
between emergence and bulbing ranged from 582 degree
days for the late planted PLK site 12 in 1986}87 to 1033 for
the early planted ELK site 38 in 1987}88 (Tables 1 and 2).
Relationships between thermal time and date of bulbing are
summarized in Fig. 2. This figure provides clear evidence for
the existence of two thresholds related to bulbing. First,
irrespective of the thermal time accumulated, bulbing did
not occur at any site before 25 Oct. This corresponds to a
day length of 135 h. It is important to note that this
photoperiod corresponds to the day length for bulbing
inferred from the median bulbing date (above). Second,
bulbing did not occur at any site before 579 degree days had
accumulated. This observation held over a range of planting
dates from 4 May to 9 Jul. The double threshold of 135 h
and 579 degree days was observed in both years and for
both cultivars.
 Lancaster et al.Photoperiod and Temperature Effects on Bulbing in Onions 427
T 4. Bulb and leaf number characteristics of Pukekohe Longkeeper (PLK) and Early Longkeeper (ELK) onions planted
on six sowing dates at Palmerston North

Thermal Estimated Estimated

Date of time emergence Mean Mean no. leaves Estimated no. leaves
600 to bulbing diameter diameter emerged no. leaves Overall no. to appear
Planting degree (date at bulbing at harvest before present at of leaves after
Site date days of bulbing) (mm) (mm) bulbing bulbing produced bulbing

PLK
1 14 May 27 Sep. 8263 (27 Oct.) 149 838 80 52 165 85
2 4 Jun. 10 Oct. 7203 (27 Oct.) 100 672 67 45 153 87
3 26 Jun. 31 Oct. 6048 (31 Oct.) 76 682 59 39 153 94
4 17 Jul. 8 Nov. 6046 (8 Nov.) 87 723 64 46 159 96
5 7 Aug. 18 Nov. 6014 (18 Nov.) 70 582 60 40 138 78
6 28 Aug. 29 Nov. 6046 (29 Nov.) 68 501 63 45 129 66
ELK
1 14 May 27 Sep. 8263 (27 Oct.) 158 725 88 54 161 73
2 4 Jun. 10 Oct. 7203 (27 Oct.) 122 656 69 46 149 80
3 26 Jun. 31 Oct. 6048 (31 Oct.) 98 613 63 41 133 71
4 17 Jul. 8 Nov. 6046 (8 Nov.) 92 559 63 43 127 65
5 7 Aug. 18 Nov. 6072 (18 Nov.) 90 547 65 45 123 58
6 28 Aug. 29 Nov. 6046 (29 Nov.) 84 467 62 46 108 46

To calculate date of bulbing at Palmerston North we used 11

the rounded figure of 600 degree days as a thermal time
No. of leaves produced after bulbing (nL)
threshold and a median of 1375 h as a photoperiod 10 4
3
threshold, (Table 3). No greater precision seemed justified,
in view of the variation expected within a population of 9
2
onions and also the slightly differing responses of PLK and 1
8 2 5
ELK. Calculations of bulbing date were made on the
following basis : bulbing cannot occur (a) if degree days are
less than 600 and day length is less than 1375 h, (b) if degree 7 1 6
3
days are less than 600 and day length is greater than 1375 h, 4
6
or (c) if degree days are greater than 600 and photoperiod
is greater than or equal to 1375 h. Thus, bulbing occurs 5
5 6
only when degree days are less than 600 and photoperiod
is greater than or equal to 1375 h (Table 3). Bulb size at the 4
estimated dates of bulbing was found to range from
68149 mm for PLK and 84158 mm for ELK (Table 4). 3
In the case of sowings 1 and 2 at Palmerston North, the 290 300 310 320 330 340
photoperiod requirement was not satisfied prior to the Day of bulbing (tB)
accumulation of 600 degree days. Therefore, in these sowings F. 3. Relationship between number of leaves appearing after bulbing
the bulbs continued to develop in diameter until the critical and day of bulbing. The relationship was determined on PLK (E) and
photoperiod for bulbing was reached. This explains the ELK (*) cultivars for sowing dates 1 to 6 (as enumerated) at
larger size at bulbing for the earlier sowings. In the later Palmerston North. Upper line, r# 089 ; lower line, r# 096.
sowing dates, the photoperiod threshold was reached prior
to the accumulation of 600 degree days and this was plants at the early sowing date had opportunity to produce
reflected in the small amount of variation in size at bulbing up to 88 leaves before bulbing, whilst those at the four later
(752085 mm for PLK and 910057 mm for ELK). planted sites only produced a mean number of 63 leaves.
The average rate of leaf emergence was 102 (005) leaves
Leaf number and bulbing per 100 degree days between emergence and bulbing. The
actual net number of leaves present at bulbing was effectively
Estimated bulbing dates in Palmerston North plantings
the same, ranging from 3952, and a median of 45,
(Table 4) were used to calculate numbers of leaves produced
irrespective of sowing date. This means that the number of
before and after bulbing from leaf appearance data. Overall
leaves which had emerged and died was greater at earlier
number of leaves produced were greatest at the earliest
planting dates.
planting date (165 for PLK ; 161 for ELK), decreased with
The number of leaves produced after bulbing varied
later sowing dates and was least at the last sowing date (129
depending on the date of bulbing and was different for ELK
for PLK ; 108 for ELK). ELK had fewer leaves than PLK
and PLK (Fig. 3). For PLK number of leaves produced
at all sowing dates. The number of leaves produced before
after bulbing (nL) was related to the day of bulbing (tB) using
bulbing was related to accumulated thermal time (Table 4,
columns 4 and 7). Bulbing terminates leaf production. Thus nL 470 (1796)310 (1139) tB00050 (000181)t#B (1)
428 Lancaster et al.Photoperiod and Temperature Effects on Bulbing in Onions
90 The time required for leaves to develop and mature as
A well as the number of leaves will influence leaf area duration
and hence the photosynthetic capacity of the plant and the
80 potential for bulb expansion. Thus size at bulbing and
number of leaves produced after bulbing are logical
Predicted size (mm)

predictors of final bulb size.

70 For Palmerston North data, bulb size at maturity (dm)
was predicted from measurement of size at bulbing (dB) and
leaf numbers to appear (nL) according to the equation :
60
dm 49 (490)21 (030) dB49 (058)nL (3)
Used predictively, this equation explained 94 % of the
50
variation in observation final bulb size (Fig. 4 A). Also, the
RMSD value relative to the mean observed value was 39 %
indicating a very precise fit. If initial bulb size alone was
40 50 60 70 80 90 used to predict size at maturity, only 476 % of the variation
Observed size (mm) was accounted for (data not shown). Therefore, size alone at
90 bulbing was not useful for predicting size at maturity,
B although early sown onions do tend to be larger and later
sown onions tend to be smaller.
80 Maturity or top down occurs when all leaves have
appeared and the neck lodges. Hence the number of leaves
yet to appear should be correlated with time to maturity. In
Predicted day

70 the Palmerston North data we found that time to maturity

60
50
50 60 70
Observed day
F. 4. Relationship between observed and predicted bulb size at
maturity (A, r# 094, RMSD 243) and maturity date (elapsed time
from bulbing) (B, r# 088, RMSD 350) for PLK (E) and ELK (*)
at Palmerston North.
This equation accounted for 89 % of observed variation.
The standard error of parameter estimates are given in
parentheses. Greater numbers of leaves appeared after
bulbing (94, 96) for PLK at sowing dates 3 and 4 than
either sowing date 1 (85) or sowing date 6 (66). Similarly
for ELK
nL 346 (290)00899 (00093)tB
This equation accounted for 96 % of observed variation.
The ELK onions had fewer leaves appearing after bulbing
than the PLK onions, and the number of leaves appearing
after bulbing decreased with later sowing date.
Prediction of bulb size and time to maturity
Variation of bulb size and time to maturity depend on the
physiological processes regulating the development of bulbs.
Bulb diameter at bulbing has been shown above (Table 3) to
be singly related to thermal time accumulation prior to
bulbing. The production of leaves after bulbing is considered
to be central to the process of bulb development since they
are key suppliers of assimilate for bulb expansion.
(tm) could be related to leaves to appear after bulbing (nL)
using :
tm 214 (580)64 (076)nL (4)
Used predictively, this equation explained 88 % of the
variation in observed time to maturity (Fig. 4 B). The
RMSD relative to the mean observed value was 51 % which
was well within the accepted range for adequate prediction.
Although a single predictive equation described the response
80 90
of both cultivars there was a distinct grouping of PLK and
ELK at either end of the maturity range.
Prediction of bulb size at maturity at Pukekohe
Used together the eqns (1)(3) constitute a model that can
be used to predict size at maturity. The predictive value of
this model can be tested for the Pukekohe sites. Numbers of
leaves to appear after bulbing at Pukekohe were calculated
using eqn (1) for PLK and eqn (2) for ELK. These data were
combined with sizes calculated at bulbing (Tables 1 and 2)
and used in eqn (3) to give predictors of bulb size at
maturity.
(2) Predicted data on bulb size at maturity are compared with
observed data in Fig. 5. Deviation around the 1 : 1 line could
be explained by variation in grower management practices
and micrometerological differences between the sites, es-
pecially with respect to temperature. The root mean square
deviation was 132 % of the mean observed value. This was
not greatly in excess of an accepted value of 10 % which
commonly defines an adequate fit between observed and
predicted data. It should also be noted that prediction of
bulb size at maturity was made at the bulbing stage when
less than 1020 % of bulb expansion had occurred.
We were unable to test the model for maturity date
predictions with an independent data set as onions were
often lifted at commercial sites before physiological maturity
could be ascertained.
 Lancaster et al.Photoperiod and Temperature Effects on Bulbing in Onions
90
80
70
Predicted (mm)
60
50
40
30 40 50 60 70 80 90
Observed (mm)
F. 5. Predicted and observed bulb size at Pukekohe for PLK (E) and
ELK (*). Bulbing date could not be computed for two sites, and a
realistic leaf number at bulbing could not be computed for PLK at one
site because of extreme lateness of sowing ; these data points were
omitted. RMSD 86.
DISCUSSION
Criterion for bulbing
In this study, we used repeated measurements of bulb and
neck diameter and used a pragmatically derived ratio of 12
as an indicator of bulbing. This non-destructive measure-
ment enabled the same plants to be used throughout the
study. Previous reports on time of bulbing have used a bulb
to neck ratio greater than 2 (Steer, 1980 a ; Mondal et al.,
1986), however, by this stage bulbing is well underway. The
cessation of appearance of new leaf blades has also been
used as an indicator of bulbing (Terabun, 1971), and is
estimated by graphs of leaf number against time. The
transition from leaf blade to bulb scale production at the
shoot apex has also been used (Holdsworth and Heath,
1950) but as with methods using leaf counts, can be
destructive.
Dual threshold model for bulbing
It has long been accepted that day length and temperature
play crucial roles in the formation and final size of the bulb.
The results reported above for field-grown PLK and ELK
onions show that in order to initiate bulbing, plants must
attain a double threshold of minimum day length of
13514 h and accumulated thermal time from emergence of
approximately 600 degree days. Bulbing is not initiated until
both these requirements are met (Table 3).
The photoperiodic requirement can be interpreted in
classical terms : onions are known to be long day plants. The
thermal time requirement is suggestive of a minimum size
requirement for bulbing. This is compatible with the
proposal of Brewster, Salter and Darby (1977) that plants
needed to have initiated a certain number of leaves to be
eligible to bulb when the day length appropriate to that
cultivar is reached. At a rate of leaf appearance of one leaf
429
per 100 degree days an accumulated thermal time threshold
of 600 degree days translates into a minimum requirement
of six emerged leaves at bulbing (Table 4). If onions
responded solely to the two thresholds then it follows that
all results would lie along the two axes of Fig. 2. However,
a scatter is observed above the two axes. At least part of this
scatter can be attributed to the fact that thermal time
records were based on data from a single meteorological site
so that calculated thermal times would overestimate the
true thermal time for cold sites and would underestimate
the true thermal time for warm sites. However, part of the
scatter may also arise because of intra-site variation and
part because the model is too simple and does not account
for interactions such as between photoperiod and tem-
perature (Steer, 1980 a).
Range of diameters at bulbing
It follows from the dual-threshold model that plants may
accumulate in excess of 600 degree day thermal time before
the critical photoperiod is reached. Thus, the early planting
of onions at Palmerston North accumulated up to 884
degree days before the critical photoperiod of 1375 h was
reached. These plants, therefore, had larger diameters and
more leaves at bulbing. It also follows from the dual-
threshold model that the critical photoperiod may be
reached before the minimum thermal time has accumulated.
In this case bulbing is delayed until the minimum thermal
time is reached. This effect was seen in late plantings at both
Palmerston North and Pukekohe and the plants concerned
have small bulb diameters and few leaves at bulbing.
Cultiar differences
The models proposed in this paper go some way to
explaining the observed difference between PLK and ELK.
These cultivars are closely related ; ELK is, in fact, a
selection from PLK (Grant, 1983). ELK is considered to be
early maturing and lower yielding. The results in this paper
suggest that ELK has a lower photoperiod threshold than
PLK (135 h s. 1375 h). Despite this difference, ELK also
has greater number of leaves produced at bulbing, fewer
total leaves produced and hence, fewer leaves produced
after bulbing. This combination explains why ELK bulbs
are larger at bulbing, smaller at maturity and earlier in
reaching maturity than PLK bulbs.
Modelling bulb size and maturity
The dual-threshold relationship of Fig. 2 and eqns (1)(4)
constitute a simple but effective model for predicting final
bulb size and maturity time. This is a top-down model in
the sense that it attempts to explain observed data without
recourse to sub-models of physiological processes. In
contrast, the model of de Visser (1994) is based on a
bottom-up synthesis of physiological concepts but is
perhaps less useful for practical applications. Despite its
simplicity, our model does offer some new insight into the
physiology of onion growth and development. Use of
430 Lancaster et al.Photoperiod and Temperature Effects on Bulbing in Onions
bulb : neck ratio of 12 rather than the more commonly
defined 20 focuses attention on physiological processes
early in the life cycle of the plant. The dual-threshold model
that emerges offers an explanation for some of the observed
results in experiments. For example, the range of size at
bulbing and synchrony of bulbing observed in sowing date
experiments can be explained in terms of satisfaction of the
thermal time threshold before the critical photoperiod is
reached. It also highlights the importance of the number of
leaves yet to appear after bulbing in determining final bulb
size and maturity. Clearly further experimentation would be
required to explore the validity and implications of this
model more fully.
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