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Anthesis, anther dehiscence, pistil receptivity and fruit development in the

Longum group of Capsicum annuum

Article  in  Animal Production Science · August 2000

DOI: 10.1071/EA99038

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C S I R O P U B L I S H I N G

Australian Journal
of Experimental Agriculture
Volume 40, 2000
© CSIRO Australia 2000

… a journal publishing papers (in the soil, plant and animal sciences)
at the cutting edge of applied agricultural research

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 Australian Journal of Experimental Agriculture, 2000, 40, 755–762

Anthesis, anther dehiscence, pistil receptivity and fruit

development in the Longum group of Capsicum annuum

M. Aleemullah, A. M. Haigh and P. HolfordA

Centre for Horticulture and Plant Sciences, University of Western Sydney Hawkesbury, Richmond,
NSW 2753, Australia.
AAuthor for correspondence; e-mail: p.holford@uws.edu.au

Abstract. Little attention has been paid to the events associated with fruit production such as flowering and
seed set in chilli peppers despite this species being a major horticultural crop. To address this deficit, we
examined flowering phenology in the cv. Red Hot Glory, a representative of the Longum group of peppers.
Measurements of bud length and width showed that these characteristics can provide an effective index of
the number of days required by a bud to reach anthesis. Anthesis itself mainly occurred during the morning
with a second, smaller peak of flower opening in the afternoon. Measurements made in spring and summer
suggested that the daily start of anthesis is controlled by daylength. In flowers that opened before 1700 hours,
anther dehiscence took place 1 h after the buds opened; in flowers that opened later, dehiscence was delayed
until the following morning. This pattern of anther dehiscence suggests that this event is partially controlled
by the flower’s endogenous rhythms. Controlled pollinations showed that the period of female receptivity,
as judged by seed set, lasted from 5 days before anthesis to 3 days after anthesis with maximum fertility
occurring on the day of anthesis. Therefore, bud pollinations are possible, and, for maximum seed set,
should be made as close to the day of anthesis as possible. This study has provided baseline data that will
aid the production of new, hybrid cultivars and will facilitate further studies on the factors affecting fertility
in this species.

Additional keywords: Capsicum annuum, chilli peppers, flowering, reproductive biology, fruit development.

Introduction interactions between the crop and the environment, the

A plant’s reproductive capacity can be one of the most
important determinants of productivity. Failure to achieve
maximum seed set not only affects yield in those crops
where the reproductive parts are harvested but can also
reduce the efficiency of plant breeding programs. The
efficiency of breeding programs and the production of
many hybrids relies on the manipulation of processes such
as pollination and pollen–pistil interactions. As Capsicum
is capable of self-pollination (Rylski 1986), production of
hybrids involves prevention of self-fertilisation by
emasculation at anthesis. As there are only a few hours
between anthesis and dehiscence (Erwin 1932) bud
pollinations would aid hybrid production. However, it is
unknown whether pollination can be successfully
conducted before anthesis.
Flower opening, anther dehiscence, and stigma
receptivity are important events that determine the success
of pollination. In turn, these processes depend on the
availability of pollen vectors, and recognition events in the
stigma and style. The timing of these events varies
between species and cultivars and it is the proper
coordination of these events that governs fruit set, yield,
and quality in many species. A detailed understanding of
the relationships between these flowering events is needed
if losses are to be prevented when environmental stresses
perturb flowering.
A study in Australia (Sumardi 1993) showed that the
longevity of the female gametophyte in the Longum group
of Capsicum annuum (chilli) is short lived. Sumardi
(1993) also demonstrated that water stress resulted in an
increase in the time required for the pollen tube to reach
the ovules, and suggested that the degeneration of the
ovule during this time led to the failure of fertilisation and
lower yields. Despite this study, basic information
concerning anthesis, anther dehiscence and the length of
time during which the female gametophyte is receptive in

© CSIRO 2000 10.1071/EA99038

756
the Longum group is lacking, as most studies on
capsicums have concentrated on the Grossum group of
C. annuum (Rylski 1986; Bakker 1989; Khan and Passam
1992; Marcelis and Baan Hofman-Eijer 1997; Pressman
et al. 1998) or on C. frutescens (Cochran and Dempsey
1966; Dempsey 1966). Hence, this investigation was
undertaken to determine the order and duration of the
events that occur during flowering and the effect of pre-
and post-anthesis pollinations on fruit development in the
chilli cv. Red Hot Glory, a representative of the Longum
group.
Materials and methods
Plant material and growth conditions
These experiments were conducted in a multispan polyhouse at
the Centre for Horticulture and Plant Sciences, University of
Western Sydney Hawkesbury, Australia, during spring 1994 and
1995 and summer 1995 using plants of the Longum group of
Capsicum annuum, cv. ‘Red Hot Glory’ (Henderson Seeds, Vic.).
The polyhouse had limited climatic controls that kept temperatures
between a maximum of 36°C and a minimum of 18°C every day but
did not provide constant temperature. The maximum and minimum
temperatures were achieved each day of the 3 experimental periods
and plants were grown under natural light conditions.
For each measurement period (spring 1994, spring 1995 and
summer 1995), 24 10-week-old seedlings, grown in cell trays, were
individually transplanted into 25 L plastic pots containing 16.5 kg
(dry wt) of potting mix. The composition of the potting mix by
volume was 3 parts composted hardwood sawdust, 4 parts
composted pine bark and 3 parts dry washed coarse sand. To
minimise evaporation from the potting mix, each pot was mulched
with 1 kg of crushed quartz gravel. The following nutrients (mg/pot)
were applied monthly: N, 537; P, 83; K, 430; Fe, 0.716; Mn, 0.376;
Zn, 0.134; Cu, 0.376; B, 0.197; Mo, 0.004. The pots were watered to
field capacity daily.
Anthesis and anther dehiscence
Three weeks after the start of flowering (14 weeks after
transplanting), when the number of flowers per plant remained
constant, observations were carried out in respect of anthesis, anther
dehiscence, pistil receptivity, bud size and the effect of pre- and post-
anthesis pollination on fruit development. Observations on anthesis
and anther dehiscence commenced before dawn and were made at
hourly intervals until after sunset, with 1100 to 1400 flowers being
counted each day. These observations were made from August to
September (early spring) in 1994 and 1995 and for a period of 4 days
in January 1995 (mid summer). Anthesis was deemed to have
occurred when the petal segments separated from each other. All
flowers that opened were recorded and the counted flowers were
pinched off to avoid recounting. At hourly intervals, 20 or more
flowers that had opened were tagged and retained on the plants to
record the time of anther dehiscence and then removed from the
plant. Anther dehiscence, in this study, was taken as the time at which
a rupture in the anther lobe was observed.
Pollination and fruit development
To collect pollen, anthers were selected at random from freshly
opened flowers of a number of plants, placed in a Petri dish and
exposed for at least 1 h to the conditions within the polyhouse to
M. Aleemullah et al.
liberate the pollen. The rubber head of a pencil was used to collect
pollen from the Petri dish and for pollinating the stigmas.
Once the observations on anthesis and dehiscence had been
completed, flowers were selected at random from each of the
24 plants and hand-pollinated at different stages of bud or flower
development to determine the period of receptivity. Pollination
commenced 7 days before anthesis (days –7 to –1) to 4 days after
anthesis (days +1 to +4). Pre-anthesis flower bud stages were
determined on the basis of bud length and width. These
measurements were recorded daily from 20 buds and the relationship
with the number of days to anthesis determined. Following the
removal of petals, flowers in the pre-anthesis stages were
emasculated, hand-pollinated, tagged and covered with a pollination
bag. Flowers to be pollinated post-anthesis were emasculated at
anthesis and covered with a pollination bag to prevent chance
pollinations. On the day of pollination, the bag was removed and
pollen was transferred to the plant’s stigma after which the flowers
were tagged and rebagged. To determine the precise time at which
the pistil ceased to be receptive, during spring, flowers were
pollinated at 2-hourly intervals, as described above, from the night of
the second day after anthesis (+2) to the morning of the fourth day
(+4). The pollination bags were removed from all flowers 6 days after
pollination to allow fruit development to occur. Receptivity was
determined on the basis of fruit set. The fruit were harvested at
maturity when the whole fruit had turned red. Fruit length, weight,
diameter (at the widest point), and seed number were determined for
35 fruit from each day of pollination except for day +3, when only
25 fruit were available.
Statistical analysis
Regression analyses and analyses of variance were performed
using either TableCurve 2D Version 4.0 (Jandel Scientific) or
COSTAT Version 4.02 (CoHort Software). Where data were not
homoscedastic according to Bartlett’s test, natural logarithm or
square root transformations were performed. Where appropriate,
multiple comparisons were performed using Student-Newman-
Keuls’ tests. Comparison of regressions was performed using the
method of Mead and Curnow (1983). A contingency chi-square test
was performed on percentage fruit set data to determine
homogeneity with 95% confidence intervals calculated according to
Snee (1974).
Results
Anthesis and dehiscence of anthers
Anthesis started before dawn (about 5% of openings)
with a large peak of opening occurring in the morning
(about 85%) and a smaller second peak in the afternoon
(Fig. 1). This pattern was similar between years and
seasons. However, in summer, the commencement of
flower opening and the period with the highest percentage
of openings were about 1 h earlier than in early spring; this
coincided with a 1 h 20 min shift in sunrise between
seasons. The time, size, and duration of the afternoon peak
did not vary between seasons despite a 3 h difference in
daylength. Once anthesis had occurred, flowers remained
open for 4 days if they were not pollinated or for 3 days if
pollinated on the day of anthesis. However, in the latter
case, petals progressively wilted from the second day after
Development in the Longum group of Capsicum annuum
pollination. Also, on average, there were 25% more
flowers (P<0.001) in summer compared to spring, with
60 and 48 flowers per plant, respectively.
Anther dehiscence and the liberation of pollen grains
occurred by the longitudinal splitting of the anthers, with
dehiscence commencing 1 h after flower opening. During
the morning period of flower opening, the petals had fully
expanded before anther dehiscence took place. However,
on days of low light intensity, dehiscence occurred without
full petal expansion. With flowers that opened in the
afternoon, dehiscence occurred although the petals did not
fully expand until the following day. Also, for flowers where
anthesis occurred at about 1700 hours, dehiscence did not
take place until the following morning, 1 h after sunrise.
Relationship between bud size and days to anthesis
Measurements of bud size were made for 7 days before
anthesis (Fig. 2). Both the length and breadth of the buds
showed a positive relationship with days to anthesis with
Spring photoperiod
Summer photoperiod
25
20
15
Anthesis (%)
10
0
0 4 8 12 16 20 24
Time of day (hours)
Figure 1. The daily pattern of anthesis in flowers of Capsicum
annuum, cv. Red Hot Glory, in early spring () and summer () in
a climate-controlled polyhouse. No significant difference was found
between the 2 sets of spring patterns, therefore, the data for the
2 years were combined. Spring data are the means of 50 days and
summer the mean of 4 days of observations. Error bars represent the
standard error of the means and where no error bar is visible, the
standard error is less than the size of the symbol. The horizontal bars
above the figure show the photoperiod in each period of observation
(average spring and summer daylengths were 11 h 34 min and 14 h
20 min, respectively).
757
these relationships having r2 values of 0.99 and 0.98,
respectively (P<0.001 in both cases). A linear relationship
existed between days before anthesis and bud length,
whereas the association between days before anthesis and
bud width was best described by a second order
polynomial equation. Measurements of bud size made on
each day before anthesis showed little variability, either
within or between plants and an analysis of variance, in
conjunction with Student-Newman-Keuls’ test, showed
that the means of both fruit length and diameter were
distinct on each day (P<0.001).
Pistil receptivity
Fruit set occurred when the gynoecium was pollinated
from 5 days before anthesis to 3 days after anthesis
(Table 1). Therefore, receptivity lasted for 9 days. The
gynoecium was most receptive on days –1, 0 and +1, as
shown by a maximum fruit set of 78–92% (Table 1). In the
pre-anthesis period, the percentage fruit set increased from
day –5 to day –1 before anthesis (Table 1). In the post-
anthesis period, receptivity decreased from day +1 to day
+3 and on the fourth day (+4) the emasculated and bagged
flowers could not be pollinated as they abscised from the
8
6
Bud size (mm)
2
0
-7 -6 -5 -4 -3 -2 -1 0
No. of days before anthesis
Figure 2. Relationship between bud length (), and bud width (),
with time before anthesis in flowers of Capsicum annuum, cv. Red
Hot Glory. Data are the means of 20 and 22 buds, respectively, for
length and width. In all cases, the error bars are less than the size of
the symbol. The equations of the lines are:
bud length (y) v. days before anthesis (x):
y = 9.13 + 0.98x (r2 = 0.99)
bud width (y) v. days before anthesis (x):
y = 3.71 – 0.09x – 0.08x2 (r2 = 0.98).
758 M. Aleemullah et al.

plant when touched. To determine the precise timing of subsets of data containing up to 90 seeds also showed that
loss of receptivity, up to 50 flowers were pollinated every there were different relationships between these
2 h, starting from 2200 hours on the night of the second parameters before and after the day of anthesis (P<0.001).
day after anthesis (+2) during spring. These pollinations The equations predicted a larger minimum fruit weight
showed that pistil receptivity decreased steadily until before than after anthesis and showed that after anthesis
1200 hours on day +3 after which time no further there was a more rapid change in fruit weight with
successful pollinations could be made (Table 2). increasing seed number than before.
Influence of pollination time on fruit characteristics
Discussion
The fruit resulting from pollinations at different pre-
In this study, we have looked at some of the events
and post-anthesis periods showed a distinct variation in
associated with pollination and fruit set in chilli peppers of
size. Fruit length, diameter, and weight as well as the
the Longum group, including the effects of pre- and post-
number of seeds per fruit increased from a minimum on
anthesis fertilisations. During the pre-anthesis period,
day –5, reached a maximum on day 0 (the day of anthesis)
good relationships were found between measurements of
then declined. The association between these parameters
bud size and the number of days to anthesis. Between each
and pollination day can be described by third order
24 h period before anthesis, sufficient differences were
polynomial equations with r2 values of 0.83, 0.55, 0.80
detected in bud length and width to allow these attributes
and 0.75, respectively (P<0.001 in all cases). The
to be used as indices of the number of days required by a
relationships between pollination day with fruit weight
bud to reach anthesis. Of these 2 measurements, length
and seed number are illustrated in Figure 3.
may provide the better index, as there was a greater
Fruit length, diameter, and weight were related to seed
absolute variation in this attribute. Therefore, it should be
number (Fig. 4). As seed number increased, fruit weight,
possible to use bud size to determine the time to anthesis
length and diameter all initially increased steeply and then
started to plateau. The relationship between fruit weight
and the number of seeds per fruit could be described by a
linear equation up to 90 seed per fruit. Past this point, there Table 2. Determination of the time post-anthesis at which pistils
was no linear relationship between seed number and fruit of C. annuum, cv. Red Hot Glory, ceased to be receptive
during spring
weight. A comparison of linear regression analyses using
Values for percentage fruit set followed by the same letter are not
significantly different from each other (P = 0.05)
Table 1. Fruit set in C. annuum following hand pollination
at different times before and after pollination Time No. of flowers Fruit set 95% confidence
(hours) pollinated (%) interval
The flowers on day +4 after anthesis abscised from the plant when
touched with a pencil and could not be pollinated
Day +2 after anthesis
Values for percentage fruit set followed by the same letter are not
2000 50 44a 13.8
significantly different from each other (P = 0.05)
2200 50 48a 13.8
2400 50 36ab 13.3
Pollination No. of flowers Fruit set 95% confidence
Day +3 after anthesis
day pollinated (%) interval
0200 50 36ab 13.3
0400 50 30ab 12.7
Pre-anthesis period
0600 50 20ab 11.1
–7 42 0a 0
0800 50 20ab 11.1
–6 56 0a 0
1000 50 16b 10.2
–5 145 26.2b 7.2
1200 50 12b 9.0
–4 84 46.4c 10.7
1400 50 0c 0
–3 94 46.8c 10.1
1600 50 0c 0
–2 85 50.6c 10.6
1800 50 0c 0
–1 64 78.1d 10.1
2000 50 0c 0
At anthesis
2200 26 0c 0
0 95 91.6d 5.6
2400 21 0c 0
Post-anthesis period
+1 60 83.3d 9.4 Day +4 after anthesis
+2 74 48.7c 11.4 0200 12 0c 0
+3 128 19.5b 6.9 0400 3 0c 0
+4 0 0a 0 0600 0 0c 0
Development in the Longum group of Capsicum annuum 759

120 12 140

100 10 120

Fruit length (mm)

Number of seeds

Fruit weight (g)

80 8 100

60 6 80

40 4 60

20 2 40

0 0 20
-5 -4 -3 -2 -1 0 1 2 3 16
Day of pollination

14
Figure 3. Relationship between seed number (), and fruit weight
(), with the day of pollination for fruit from the Longum group of
Fruit diameter (mm)
C. annuum. 12

10
so facilitating bud pollination and the production of hybrid
chilli cultivars.
A study of anthesis, using several unnamed varieties of 8
peppers, suggested that the duration of anthesis is similar
in all capsicums with only 1 period of opening that is 6
completed within 5 h of sunrise (Erwin 1932). Quagliotti
(1979) also suggested that there is only 1 period of
14
anthesis in capsicums, which is complete by 0900 hours.
In contrast, this study with the cv. Red Hot Glory showed 12
that anthesis occurred in 2 main peaks each day: a
10
Fruit weight (g)

morning peak that lasted from pre-dawn to 1200 hours and

a second peak in the afternoon (1400–1500 hours). The 8
detection of the afternoon peak may be due to the large
number (more than 1100 per day) of flowers observed. 6
Therefore, this study shows that the pattern of anthesis
4
within the peppers is not as similar as Erwin’s (1932) study
suggested, an idea reiterated in Rylski’s (1986) review. 2
In his mid-summer study, Erwin (1932) found that
flowers did not open before dawn and that 90% of flowers 0
had opened within 2.5 h of sunrise. In our study, we found 0 40 80 120 160 180
that 5% of flowers had opened before sunrise, 6–9 h after No. of seeds per fruit
dawn were required for 90% of flowers to open, with the
remainder opening during the afternoon peak. In addition, Figure 4. Relationships between seed number (x) and fruit length,
Erwin (1932) reported that the period during which diameter and weight of Capsicum annuum cv. Red Hot Glory. These
flowers remain open in capsicums is short, being less than relationships were best approximated by saturating, kinetic
24 h. During both summer and early spring in our study, equations of the form y = (a – b) exp (– cx) + b where a = 34.42;
once anthesis had occurred, the flowers remained open for b = 126.46; c = 0.05 (length v. seed number, r2 = 0.91), a = 8.52;
b = 14.81; c = 0.007 (fruit diameter v. seed number, r2 = 0.46) and
3–4 days depending on whether the plants were pollinated a = 0.93; b = 11.53; c = 0.017 (fruit weight v. seed number,
or not. Again, our results show that anthesis in the genus r2 = 0.80).
760
Capsicum is more variable than Erwin proposed and
additional work is required to determine if further
variation exists, other than in the Longum group.
The reason for the 2 periods of anthesis is unclear.
Capsicums, like other members of the Solanaceae, have
wet stigmata that secrete a fluid (Heslop-Harrison and
Shivanna 1977). This stigmatic fluid has a number of roles
including pollen adhesion, hydration and germination
(Hoekstra and Bruinsma 1975). Exposure to the low
humidities that result from temperature increases may
cause the stigmatic surface to dry thereby hindering pollen
germination. Therefore, during the hottest part of the day,
it may be an advantage to plants if anthesis is delayed until
temperatures drop. However, in other members of the
Solanaceae, such as Nicotiana and Petunia, the exudate
does not play an important role in pollen germination, as
young stigmata that are free from exudate support
satisfactory germination (Shivanna and Sastri 1981).
Therefore, the role of the stigmatic fluid and any effect of
the environment on the germination of capsicum pollen
need further investigation.
Whilst it is well documented that temperature
influences flower development in capsicums (Erwin 1932;
Polowick and Sawhney 1985; Rylski 1973, 1986;
Pressman et al. 1998) little is known of the effects of
daylength in peppers other than that they are short day
plants (Auchter and Harley 1924; Deats 1925; Cochran
1942). Our data suggest that flower opening in the
Longum group of C. annuum is regulated, at least in part,
by daylength. The 2 years of early spring observations
showed identical patterns of flower opening and there was
an approximate 1 h shift forwards in the morning peak
during summer as a result of a 1 h 20 min earlier sunrise.
The daily temperatures during these 3 periods of
observations were similar (due to temperature regulation
in the polyhouse), however, the light intensity would have
differed substantially between early spring and mid-
summer. It would be interesting to determine whether
other factors, such as temperature or light intensity, also
influence flowering phenology in chillies.
The timing of anther dehiscence in many species is
associated with mechanisms to ensure cross-pollination,
with final yield and fruit quality being determined by the
co-ordination of this event with stigma receptivity. In Red
Hot Glory, dehiscence occurred 1 h after anthesis ensuring
that pollen grains are liberated at the time of maximum
stigmatic receptivity. The slight delay between anthesis
and dehiscence may be required for anthers to dry
sufficiently for the cells of the stomium to separate so that
the pollen grains can be shed. Erwin (1932) reported that
the anthers of other capsicum flowers did not dehisce until
M. Aleemullah et al.
3 h after the flower petals opened which also suggests that
dehydration may affect the timing of dehiscence.
An understanding of stigmatic receptivity in crop
plants is important for several reasons. Firstly, seed set
may be maximised by pollinating receptive stigmas at the
optimal time. Secondly, pre-anthesis bud pollination may
be able to overcome self-incompatibility mechanisms,
thereby facilitating the production of hybrid cultivars.
Lastly, the period of receptivity in emasculated flowers of
self-pollinated crops may be brief as in cotton, wheat and
tomato giving a limited time when pollinations can be
performed (Leopold and Kriedemann 1975; Shivanna
et al. 1997). A number of biochemical tests are available
for the assessment of receptivity; however, controlled
pollinations, the technique used in this study, provide the
most reliable results as well as being a quantitative
procedure (Dafni 1992). As with many other species,
receptivity in Red Hot Glory commenced before flower
opening, starting on day –5 before anthesis and lasting
until day +3 after anthesis: this gives an effective
pollination period (EPP) of 9 days. The post-anthesis
period of receptivity of Red Hot Glory is similar to that of
C. frutescens where maximum receptivity occurred on the
day of anthesis and lasted for 3 or 4 days after anthesis
depending on time of the year (Cochran and Dempsey
1966).
Although receptivity in Red Hot Glory commenced
5 days before anthesis, the percentage of pollinations
resulting in fruit set varied considerably during this period,
with only 26% of flowers pollinated on day –5 producing
fruit compared with 78% on day –1. Therefore, if bud
pollinations are required, these pollinations should be
performed close to the day of anthesis. For this, a reliable
estimate of the number of days to anthesis is needed and,
as we have shown, this can be obtained from
measurements of bud size. Although this study suggests
that the EPP in the Longum group is about 9 days, only the
pollinations made on days –1, 0 and +1 resulted in a high
percentage of fruit set. Therefore, it may be better to
consider that the useful pollination period lasts for only
3 days, which supports Cochran and Dempsey’s (1966)
assertion that the duration of receptivity is short in all
capsicums.
Reliable estimates of ovule longevity, made under
optimal climatic conditions, are also required to determine
the cause of poor seed set, yields, or quality. There is
evidence from cytological studies that under certain
environmental conditions, such as frost damage, ovules
may be non-functional by the time pollen tubes arrive in
the ovary (Stosser and Anvari 1982). Receptivity is
frequently terminated by the abscission of the flower bud
Development in the Longum group of Capsicum annuum
(Leopold and Kriedemann 1975; Shivanna et al. 1997)
while in tomato, the ovules remain viable after abscission
(Leopold and Scott 1952). In our study, fruit set declined
quickly after its maximum on the day of anthesis with no
successful pollinations possible after 1200 hours on day
+3; therefore, receptivity ceased before abscission. In the
cultivar used in this study, water stress increases the time
required for the pollen tube to reach the ovules which leads
to reduced seed set (Sumardi 1993). He suggested that
ovule degeneration was the cause of this reduction in
fertilisation. However, under favourable conditions,
pollination should not be a major factor affecting yields as
fruit set did not drop significantly until day +2 after
anthesis. Therefore, explanations for poor fruit set under
these circumstances should be sought from factors
affecting the quality of gametes or their union.
Fruit size in Red Hot Glory positively correlated with
seed number as has been found in other capsicums.
However, in our study, for the fruit resulting from pre- and
post-anthesis pollinations, the overall relationship was not
linear and an asymptotic equation was required to describe
the data. In the Grossum group, a significant linear
relationship was found between seed number and fruit dry
weight; this relationship was not improved by fitting a
polynomial equation (Marcelis and Baan Hofman-Eijer
1997). Similar linear relationships have been reported by
Rylski (1973), de Ruijter et al. (1991) and Shipp et al.
(1994). In our study, linear relationships between seed
number and fruit size could only be found when regression
analyses were performed on subsets of data where the fruit
contained less than 90 seed per fruit.
In the studies performed on members of the Grossum
group by Marcelis and Baan Hofman-Eijer (1997), Rylski
(1973), de Ruijter et al. (1991) and Shipp et al. (1994),
seed number was manipulated by pollen load at the time of
maximum receptivity. In this investigation, variation in
seed number resulted from pollinations on different days
before or after anthesis. Fruits are sinks that compete for
photosynthate, other nutrients, and water required for
growth. The size and strength of each sink is determined
by its metabolic activity which, in turn, appears to be
controlled by the production of phytohormones (Coombe
1965; Leopold and Kriedemann 1975; Varga and
Bruinsma 1976; Goodwin 1978). In a number of species,
including chillies, developing seeds produce high levels of
auxin (Leopold and Kriedemann 1975). The differences
between our results and those from the other studies on the
relationship between seed number and fruit size may be a
consequence of changes in the ovular and surrounding
tissue’s ability to produce or respond to this hormone
during bud and fruit development. Evidence to support
761
this view comes from the fact that the regression equations
of subsets of fruit coming from pollinations before and
after anthesis were different.
A further difference that is related to fruit development
in the Longum and Grossum groups can be found in their
ability to produce parthenocarpic fruit. Parthenocarpic
fruit can be readily obtained in the Grossum group
(Cochran 1936; Rylski 1973; Polowick and Sawhney
1985; Shifriss and Eidelman 1986; Marcelis and Baan
Hofman-Eijer 1997). However, no reports of naturally
occurring parthenocarpy can be found in the Longum
group and in our study, all fruit contained seed. The
developmental processes that allow fruit of the Grossum
group to develop without seeds may also account for the
difference in the relationships between fruit size and seed
number found between the 2 groups.
Marcelis and Baan Hofman-Eijer (1997) only found a
linear relationship between fruit size and seed number in
half their studies. In the other half, supplementary
pollination caused no increase in fruit weight, and they
hypothesised that the effects of seed number may saturate
at high values. Our results provide direct evidence to
confirm this suggestion, as firstly, no increase in fruit size
or weight was observed once fruit contained over 90 seeds.
Secondly, the slope of the regression lines for subsets of
data containing <90 seeds per fruit became less as the
subsets contained data from fruit with higher numbers of
seeds. Thirdly, a regression analysis on a subset of data
from fruit containing >90 seeds found no relationship with
fruit size. Lastly, the relationships between seed number
and fruit size could be described by kinetic equations that
saturate before reaching an asymptote.
In this study, we have detailed flowering behaviour and
fruit set in chilli peppers from the Longum group. This
information will enable plant breeders to produce cultivars
more efficiently, as breeders seem to be unaware that pre-
anthesis pollinations are possible (Jaime Mendez pers.
comm.). This study also provides baseline data that can be
used to discover the causes of poor fruit formation during
stress conditions. In particular, as the useful pollination
period in the Longum group can be considered to be only
3 days, it would be desirable to investigate further the
events from pollination through fertilisation to embryo
development to determine the factors leading to successful
fruit set or abortion.
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Received 1 April 1999, accepted 10 March 2000