Open Agriculture.

2017; 2: 580–588

Rapid communication

Duwini Padukkage, Gamini Senanayake, Sudarshanee Geekiyanage*

Photoperiod sensitivity of very early maturing

Sri Lankan rice for flowering time and plant
architecture
https://doi.org/10.1515/opag-2017-0061
Keywords: Days to flowering, photoperiod sensitivity, very
received October 30, 2016; accepted September 22, 2017
early maturing Sri Lankan rice accessions
Abstract: Unavailability of information on photoperiod
sensitivity of traditional rice is a disadvantage in breeding
rice for adaptation to changing climatic conditions and for
optimum plant architecture. This experiment was conducted
1 Introduction
to address the above problem through determination of
Rice (Oryza sativa L.) is considered as the staple food in
variation in days to flowering (DF) and morphological
many Asian countries (Izawa 2008). Sri Lanka possesses a
traits of twenty eight selected very early maturing Sri
wide gene pool of traditional rice with high environmental
Lankan traditional rice accessions under three photoperiod
adaptability through different morphological and
conditions: short day (SD), day neutral (DN) and long day
physiological characters including plant height, days
(LD). Although rice is considered to be a facultative short
to flowering (DF), grain shape, pericarp colour and
day plant, 12 accessions (2088, 2979, 4615, 3970, 4245, 2091,
presence of awns (Team of NRC Research Project 12-129,
3450, 3883, 4217, 6741, 3738 and 3677) flowered significantly
2014, 2015). Rathnathunga et al. (2016) evaluated 384 rice
late under SD over LD, while accession 4223 flowered
accessions from 53 Sri Lankan traditional rice varieties for
significantly late under DN. Fifteen accessions (3943, 4042,
morphological and flowering time variation and found
4734, 3693, 4513, 3845, 4390, 4144, 4220, 4223, 4237, 4387,
that 39 accessions did not flower during the experimental
Bg 300, At 308 and Bg 379-2) flowered significantly early
period of 210 days in late short day season. According to
under SD over LD. Four accessions (3457, 3884, 6305 and
PGRC (1999), there were 352 accessions, which flowered
4358) were non-responsive to photoperiod for flowering
at or below 75 days under tested conditions in Sri Lanka.
time. Plant height significantly increased only in accession
Mechanisms related to the photoperiodic flowering
4217 under SD. Higher plant height under DN was obtained
in rice plants were further studied with recent molecular
from At 308 and accession 3883. Photoperiod did not affect
approaches (Izawa et al. 2003; Searl and Coupland 2004).
plant height in accession 4237 and Bg 379-2. LD only or both
In Northern Japan, experiments have been conducted
LD and DN conditions produced significantly higher plant
to develop early flowering and photoperiod insensitive
height irrespective of photoperiod responsiveness for DF in
cultivars (Izawa 2008). Extreme early flowering
rest 26 accessions. Variation in response to photoperiod for
behaviours could be observed in the northernmost regions
DF and plant architecture in very early maturing Sri Lankan
of the world as well (Uwatoko et al. 2008). Molecular
rice indicates the potential diversity of genetic factors for
and morphological studies on photoperiod insensitivity
photoperiod response for future use in rice breeding.
and early flowering nature of rice plant leading to
identification of responsible genetic factors help to breed
*Corresponding author: Sudarshanee Geekiyanage, Department new rice varieties adapted to different latitudes, altitudes,
of Agricultural Biology, Faculty of Agriculture, University of Ruhuna, seasons and changing climatic conditions (Izawa 2007;
Mapalana, Kamburupitiya, Sri Lanka, E-mail: sudarshanee@agbio. Khush 2001; Uwatoko et al. 2008; Xu et al. 2014).
ruh.ac.lk
Rice is a facultative short day plant. It exhibits a wide
Duwini Padukkage, Faculty of Graduate Studies, University of Ruhu-
na, Matara, Sri Lanka range of variation in degree of sensitivity to photoperiod,
Gamini Senanayake, Department of Agricultural Biology, Faculty of which controls its growth and flowering (Izawa 2008;
Agriculture, University of Ruhuna, Mapalana, Kamburupitiya, Sri Ogiso-Tanaka 2013; Uwatoko et al. 2008; Yano et al. 2001;
Lanka Xu et al. 2014). Sri Lankan rice varieties also exhibit a wide

Open Access. © 2017 Duwini Padukkage et al., published by De Gruyter Open. This work is licensed under the Creative Commons Attribution-
NonCommercial-NoDerivs 3.0 License.
  Photoperiod sensitivity of very early maturing Sri Lankan rice varieties 581

variation in sensitivity to photoperiod, which controls
plant growth and flowering leading to difficulties in
cultivation (Chandraratna 1964). Rice plant architecture is
affected by flowering time in response to photoperiod (Wei
et al. 2010). According to Team of NRC Research Project
12-129 (2015), different rice varieties differed markedly in
flowering initiation during the short day season of year
2013-2014, where several very early maturing accessions
were recorded.
Despite potential advantages of traditional rice with
nutritional and medicinal properties, Sri Lankan rice
cultivation is confined to new improved varieties since
the last few decades (Central Bank, 2014). Although
Sri Lanka had already reached self-sufficiency in rice
production, reduction of almost 1 million tons of rice was
recorded in 2014 due to severe drought. Adverse climatic
and environmental conditions continued in 2016 and 2017
and rice production in 2017 is forecast at 40% reduction in
comparison to previous year (FAO of the UN 2017). Such
effects could reduce the rice production of the country in
the future. Therefore, increasing rice production through
breeding rice varieties with very early maturity and
photoperiod insensitivity is a current necessity to deal
with unfavourable climatic changes.
Field data on flowering time variation within season
and sensitivity to photoperiodic season is a knowledge
gap for utilizing Sri Lankan traditional rice varieties for
farmer introductions and conventional and molecular
breeding programmes. Identification of early maturing
rice accessions with photoperiod insensitivity was
aimed in this study, where selected very early maturing
accessions were tested for their photoperiodic responses
for flowering initiation and morphological traits under
controlled photoperiod conditions. Identification and
modulation of genetic factors and developing early-
flowering photoperiod-insensitive varieties would be
useful for higher yield and optimum plant structure
manipulation, which would ultimately lead to the global
food security in the era of climate change.
2 Materials and Methods
2.1 Plant material selection
Twenty eight accessions (of 70 - 75 DF) were chosen
(Table 1) from Team of NRC Research Project 12-129 (2015).
Three improved rice varieties of Bg 300, Bg 379-2 and At
308 were included as control.
2.2 Site description
The experimental site for photoperiodic experiment was
located in Department of Agricultural Biology, Faculty
of Agriculture, University of Ruhuna (6°17’0” of North
and 81°17’0” of East, 24 m above mean sea level). The
experiment was conducted in a photoperiodic chamber
(where SD, LD and DN conditions were imposed) from July,
2014 to January, 2015. Average monthly temperatures of

Table 1: Selected traditional rice accessions

No. Accession Variety name No. Accession Variety name

1 2088 Sudu heenati 15 4144 Mudukiri al

2 2091 Sudu heenati 16 4217 Ballawala

3 2979 Heenmurunga 17 4220 Batapolayal

4 3450 Andaragaha wee 18 4223 Dhemas wee

5 3457 Ittikulama 19 4237 Hathe pas davase wee

6 3677 Herathbanda 20 4245 Iri wee

7 3693 Herathbanda 21 4358 Sudumada al

8 3738 Mudukiri al 22 4387 Baana wee

9 3845 Kahamalan 23 4390 Chellanayagam

10 3883 Japan heenati 24 4513 Galu wee

11 3884 Japan heenati 25 4615 Driver wee

12 3943 Cheenadi 26 4734 Inthiankaruppa

13 3970 Mudukiri al 27 6305 Enawakka

14 4042 Gam podi wee 28 6741 Kahasinanayam
582 D. Padukkage, et al.
29.50C in July, 28.60C in August, 29.20C in September, 28.50C
in October, 28.40C in November, 28.10C in December, 2014
and 290C in January, 2015 were recorded.
2.3 Experimental design and management
practices
Nursery grown seedlings were maintained for 14
days under natural photoperiod, that prevailed until
transferring to the mud pots of 20 cm diameter. Plants
were grown in three photoperiod conditions according in
a Completely Randomized Design (CRD) in 3 replicates:
short day (SD) condition with 8 hours of light and 16
hours of darkness, day neutral (DN) condition with 12
hours of light and 12 hours of darkness and long day (LD)
condition with 14 hours of light and 10 hours of darkness.
All plants were exposed to 8 hours of natural light while
the plants in DN and LD chambers were exposed to light
from white florescent bulbs. Fertilizer was added at 2,6
and 10 weeks after plant establishment, according to the
recommendation of Department of Agriculture, Sri Lanka:
Muriate of Potash (MOP) 0.157 g per plant and Triple Super
Phosphate (TSP) 0.196 g per plant and Urea 0.157 g per
plant in a pot (as Urea 50 kg/ha, TSP 62.5 kg/ha, MOP 50
kg/ha) were applied. Plants were watered regularly.
2.4 Plant measurements and data analysis
Data was taken at flowering stage. Days to flowering (DF),
plant height (PH) and culm number (CN) were measured
according to the guidelines of Standard Evaluation System
for Rice (PGRC 1999). Two-way analyses of variance
(ANOVA) and analysis of means (ANOM) were conducted
using Minitab® (Version 15.1.0.0.) statistical software
and p < 0.05 was used to determine significance. Mean
separation was carried out through Duncan Multiple
Range Test (DMRT) using SAS software (9.1 version, USA)
Table 2: Variation of days to flowering, plant height and tiller number under three photoperiods
Photoperiod Min / Max Days to flowering
condition
Long-day Min 60 (Accession 2088)
Max 113 (Accession 4144)
Day-neutral Min 60 (Accession 2088)
Max 107 (Accession 4387)
Short-day Min 68 (Accession 2088)
Max 110 (Accession 3970)
to determine the significant differences among accessions
for selected traits.
3 Results
3.1 Variation in morphological traits among
photoperiods
Variation of DF, PH and CN of accessions among three
photoperiods is given in Table 2.
Long-day treatment: The minimum value for DF was
observed in accession 2088 (Sudu heenati) having DF of
60 ± 0.706 days (CV % = 1.66%) and maximum DF of 113
± 1.767 days (CV % = 2.22%) was given by accession 4144
(Mudukiri al): The lowest and highest PH were recorded
from improved rice variety At 308 (88.7± 8.96 cm; CV %
= 17.5%) and accession 6305 of variety Enawakka (197.0 ±
0.707 cm; CV % = 0.51%) respectively. The CN varied in LD
condition giving the lowest value by accession 2088 (2 ±
0.0, CV % = 0.0%) and the highest value as 10 ± 1.06 (CV %
= 15.78%) by accession 3970 of Mudukiri al.
Day-neutral treatment: Accession 2088 exhibited the
lowest DF of 60 ± 0.0 days (CV % = 0.0%) while accession
4387 (Baana wee) exhibited the highest DF of 107 ± 3.68
days (CV % = 5.95%). The lowest and highest PH were
recorded in Bg 379-2 (93 ± 3.53 cm; CV % = 5.37%) and in
accession 6305 (184.5 ± 2.47; CV % = 1.89%) respectively.
Accession 4513 (Galu wee) exhibited the lowest CN as 2
± 0.353 (CV % = 33.3%) and accession 3884 (Japan wee)
exhibited the highest CN as 7 ± 2.49 (CV % = 3.63%).
Short-day treatment: Accession 2088 exhibited a
similar response to LD and DN in SD also giving the lowest
DF of 68 ± 0.35 days (CV % = 0.74%). Accession 3970
(Mudukiri al) recorded the highest DF in SD condition (110
± 0.707, CV % = 0.909%). The minimum and maximum PH
were given by accession 3943 (Cheenadi), 67.5 ± 2.82 cm
(CV %= 3.22%) and accession 6305 of Enawakka (134.5 ±
9.54, CV %= 10.03%) respectively. The lowest CN was given
Plant height (cm) Tiller number
88.7 (At 308) 2 (Accession 2088)
197 (Accession 6305) 10 (Accession 3970)
93 (Bg 379-2) 2 (Accession 4513)
184.5 (Accession 6305) 7 (Accession 3884)
67.5 (Accession 3943) 2 (Accession 3677)
134.5 (Accession 6305) 12 (Accession 3884)
  Photoperiod sensitivity of very early maturing Sri Lankan rice varieties
by accession 3677 of Herathbanda (2 ± 2.82, CV %= 4.65%)
and the highest CN was given by accession 3884 of Japan
heenati (12 ± 3.74, CV %= 6.05%).
3.2 Effect of photoperiod on days to
flowering
Twelve accessions flowered significantly late under SD
condition over LD condition or DN condition: Six flowering
time responses were identified among the 12 accessions.
Accessions 2088 (Sudu heenati), 2979 (Heenmurunga),
4615 (Driver wee), 3970 (Mudukiri al) and 4245 (Iri wee)
flowered significantly early under DN and LD. Accession
2091 flowered significantly early only under LD, while DF
under SD and DN were not different. Accession 3450 also
flowered significantly early under LD, while DF under
SD was greater than that of DN. Meanwhile, accession
3883 flowered also early under LD and DF under DN was
greater than that of SD. Accessions 4217 and 6741 flowered
earlier under LD over SD, while flowering times under
SD and LD were not significantly different to that of DN.
Significantly early flowering under DN only was recorded
from accessions 3738 and 3677, where DF under SD and LD
were not significantly different (Figure 1).
Fifteen accessions flowered significantly early under SD
over LD or DN, which exhibited 6 responses to photoperiod:
accession 3943 flowered significantly early under SD, while
DF under DN was not different to those of SD and LD. DF
under DN was greater than that under LD in accession 4042
and improved rice Bg 379-2. DF under LD was greater than
those of SD and DN in 4734. DF under LD and DN were not
different in accessions 3693 and 4513. DN condition resulted
in significantly late flowering over SD and LD in accessions
3845 and 4390. DF under LD was greater than that of DN,
while DF under SD was the earliest in accessions 4144,
4220, 4237 and 4387, and improved rice Bg 300 and At 308.
Accession 4223 flowered significantly late under DN.
Four accessions (3457, 3884, 4358 and 6305) did not
respond to photoperiod as DF of above accessions were
not significantly different among photoperiods (Figure 1).
3.3 Effect of photoperiod on culm number
and plant height
Culm number varied from 2 - 12 among accessions in
3 photoperiods, which did not exhibit a relationship
between photoperiods (Figure 2).
Plant height was affected by photoperiod in different
accessions (Figure 3): Except for accession 4237 and
583
improved rice Bg 379-2, other 29 accessions responded
to photoperiod for plant height. Only accession 4217
produced significantly higher plant height under SD.
Accession 3883, and improved rice At 308 produced
significantly higher plant height only under DN. Other
accessions produced significantly higher plant heights in
LD only or LD and DN irrespective of flowering response
in LD and DN.
4 Discussion
More than 70 rice genes known to be associated with
flowering time had been detected through mutants,
transgenic plants and natural variation (Hori et al.
2016). Rathnathunga et al. (2016) developed a mini core
collection of Sri Lankan traditional rice based on wide
naturally occurring flowering time variation. In this
study, a wide variation in flowering time for photoperiod
was detected among selected early maturing accessions:
twelve accessions were identified as delayed flowering
under inductive SD condition, while fifteen accessions
were early flowering under SD condition. Four accessions
did not respond to photoperiod for flowering. Even
though new improved rice varieties grown in Sri Lanka
are known to be photoperiod-insensitive during 2 mild
photoperiod seasons of Sri Lanka, Bg 300, Bg 379-2 and
At 308 were early flowering under tested extreme SD
condition. Accessions 3970 and 4144 of variety Mudukiri al
exhibited different responses to photoperiod for flowering
initiation as accession 3970 was late flowering under SD,
while accession 4144 was early flowering under SD and
late flowering under LD: Rathnathunga and Geekiyanage
(2017) reported that days to reach fifth leaf stage, which is
an indication of length of vegetative phase, in accessions
3970 and 4144 were significantly higher under SD and LD
respectively suggesting that different effect of photoperiod
on vegetative phase may lead to flowering time variation
between accessions of the same variety. According to
Hori et al. (2016), previous researchers had indicated that
allelic variation of Hd1, Ghd7, DTH8, Hd6, OsPRR 37, DTH2
and Ehd4 among different genotypes led to the regional
adaptation through flowering time variation. Takahashi
et al. (2009) reported the presence of non-functional Hd1
allele in Sri Lankan traditional rice variety Vandanan
leading to short day insensitivity. The development of
improved varieties with insensitivity to photoperiod by
introduction of the photoperiod-insensitive genes was
done during past few decades and these varieties can
be grown during any season and in most of the tropical
and subtropical countries. These improved varieties
584 D. Padukkage, et al.

Figure 1: Effect of photoperiod on a variety of days to flowering responses among Sri Lankan very early maturing rice accessions: 1) Twelve
accessions flowered significantly late under SD condition over LD condition or DN condition; 2) Four accessions didn’t respond to photope-
riod for flowering time; 3) Fifteen accessions flowered significantly early under SD over LD
  Photoperiod sensitivity of very early maturing Sri Lankan rice varieties 585

Figure 2: Effect of photoperiod on culm number variation among Sri Lankan very early maturing rice accessions: 1) that flowered early under
LD condition; 2) that flowered early under DN condition; 3) that flowered early under SD condition
586 D. Padukkage, et al.

Figure 3: Effect of photoperiod on plant height variation among Sri Lankan very early maturing rice accessions: 1) that flowered early under
LD condition; 2) that flowered early under DN condition; 3) that flowered early under SD condition
  Photoperiod sensitivity of very early maturing Sri Lankan rice varieties
are with wide adaptability and short vegetative growth
period, which results with higher per day productivity
(Khush 2001; Xu et al. 2014). Mutations in DTH8 cause a
weak photoperiod response and early flowering in Asian
cultivated rice. It is important in regulating the plant
height and yield potential in rice. A chromosome segment
of DTH8 has a remarkable effect on flowering time, plant
height, and yield (Wei et al. 2010). Plant height was higher
under LD in most early maturing accessions irrespective
of flowering response. Variation in tiller number did not
relate to photoperiod in tested accessions. Meanwhile,
late maturing SD sensitive Sri Lankan traditional rice
produced significantly higher number of tillers under LD
(Geekiyanage et al. 2012). Xia et al. (2012) reported that
increased tiller number and early flowering was resulted
in transgenic lines overexpressing the microRNA OsmiR
393, which could affect two rice auxin receptor genes.
Our results indicate that very early maturing Sri
Lankan traditional rice germplasm consists of three groups
of rice accessions based on response to photoperiod for
flowering time as short day-sensitive, long day-sensitive
and day-neutral groups. According to Vergara and Chang
(1985) and Chandraratne (1965) many of the reported
LD sensitive accessions were found to be SD sensitive
in subsequent testing. Therefore, the accessions that
flowered late under SD must be tested to confirm late
flowering response under SD in repeated experiments.
Identification of genetic factors responsible for a variety
of flowering responses to photoperiod is important in
rice breeding in the future: the variety of responses of Sri
Lankan traditional rice accessions to photoperiod is of
importance to the research community for identification
of genetic factors in Sri Lankan traditional rice gene pool.
The critical information provided can be used to breed
novel varieties with worldwide adaptability through
photoperiod insensitivity.
Acknowledgement: Authors acknowledge National
Research Council, Sri Lanka for funding through NRC
12-129 grant to SG and Plant Genetic Resources Centre, Sri
Lanka for provision of traditional rice seeds.
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