105
INFLUENCE OF PHOTOPERIOD AND TEMPERATURE
1
ON THE DEVELOPMENT OF WINGED BEAN PLANTS
Marlene A. Schiavinato 2 and Ivany F.M. Válio 3
Departamento de Fisiologia Vegetal, Instituto de Biologia, Universidade
Estadual de Campinas, CP 6109, Campinas, SP, 13083-970, Brasil.
ABSTRACT - Plants of winged bean (Psophocarpus
tetragonolobus D.C.) were maintained under
photoperiods of 10, 12, 13, 14, 16, 20 hours and
different day/night thermal regimes (20/15; 25/20 and
30/25oC). Flower initiation and tuberization were only
observed in plants under photoperiods below 20h.
Winged bean may be considered therefore, a short
day plant. In relation to day/night thermal regimes,
plants under a 13h photoperiod and temperatures of
20/15oC presented reduced growth. Plants under
30/25oC presented an increase in shoot dry mass and
leaf area. Nevertheless, plants under inductive
photoperiods only flowered when grown at 25/20oC
day/night temperatures. These plants also presented
the greatest tuberous root dry mass.
Additional index terms: flowering, growth,
Psophocarpus tetragonolobus , tuberous roots.
INFLUÊNCIA DO FOTOPERÍODO E DA
TEMPERATURA NO
DESENVOLVIMENTO DE PLANTAS DE
FEIJÃO ALADO
RESUMO - Plantas de feijão alado (Psophocarpus
tetragonolobus D.C.) foram mantidas sob fotoperíodos
de 10, 12, 13, 14, 16, 20 horas e diferentes regimes
de temperatura dia/noite (20/15; 25/20 e 30/25 oC). A
iniciação floral e a tuberização somente foram
observadas sob fotoperíodos inferiores a 20h.
Portanto, a espécie pode ser considerada uma planta
de dias curtos. Com relação aos regimes térmicos,
plantas cultivadas sob fotoperíodo de 13h e 20/15 oC
apresentaram crescimento reduzido. As plantas em
30/25oC apresentaram aumento de massa seca da
parte aérea e de área foliar. Todavia, plantas sob
fotoperíodos indutores somente floresceram quando
mantidas em temperaturas de 25/20oC. Estas plantas
também apresentaram a maior massa seca de raízes
tuberosas.
1
Received in 05/03/1996 and accepted in 27/05/1996.
2
Prof. Assistente Dr
3 established that temperature influences responses to
Ph.D.(Lond.), Professor Convidado Titular
R.Bras.Fisiol.Veg., 8(2):105-110, 1996.
Termos adicionais para indexação: crescimento,
floração, Psophocarpus tetragonolobus, raízes
tuberosas.
INTRODUCTION
Psophocarpus tetragonolobus (winged bean) is
considered a species well adapted to warm, humid
tropics (Noda et al, 1984), although its distribution
includes drier, colder and higher regions (Khan, 1978).
Most of the components (if not all) of any ecosystem
have their particular characteristics in response to the
photoperiods to which they are exposed (Vince-Prue,
1975). Garner & Allard (1920), clearly demonstrated
that flowering and many other plant responses may be
controlled by short or long days, depending on the
plant.
Vince-Prue (1975) quoted 23 species of plants that
show an increase in leaf area when exposed to long
days, and 3 species that present this effect under short
days. Brondum & Heins (1993) observed that lateral
shoot count and length decreased and tuberous root
mass increased as photoperiod decreased from 16 to
10 hours.
Since many storage organs are important sources
of food products, the factors that influence their
development have received considerable attention. In
several species the formation of these organs
depends on, or is accelerated by, exposition of leaves
to specific photoperiods. Frequently, tuber and
tuberous root formation are promoted by short days
(Murty & Banerjee, 1977; Okubo et al, 1992).
According to some authors, winged bean plants
require short days for flower and tuber formation
(Wong & Schwabe, 1979; Klib-Ngern & Bautista,
1982). This species rarely flowers when growing out of
the tropics. This has been attributed more to daylength
differences rather than to temperature (N.A.S., 1975).
Since in humid, tropical zones, there is a minimal
variation of daylength, the topography may be the
result of differences in temperature. It is well
photoperiod, growth, tuber development and flowering
106

of many species (Arulrajah & Ormrod, l973; Brewster,
1983; Cao & Tibbitts, 1994). For tropical species, the
evidence suggests that temperature is just as
important as daylength in influencing growth and
flowering (Herath & Ormrod, 1979; Wong & Schwabe,
1979).
The aim of this work was to study the behavior of P.
tetragonolobus plants, when submitted to different
photoperiods and day/night temperatures.
MATERIALS AND METHODS
temperatures of 20/15, 25/20 and 30/25 oC and a 13h
photoperiod.
Statistical design and analysis- In all
experiments, the layout was a randomized block
design, with 10 replicates for each treatment. Data
were subjected to an analysis of variance and a
comparison of means carried out using Tukey’s test
(Snedecor & Cochran, 1967). In experiments where
plants were harvested at more than one period, the
statistical analyses were made for each harvest
separetely.

Plant material- Seeds of Psophocarpus

tetragonolobus (L.) D.C. (winged bean) obtained
RESULTS AND DISCUSSION
originally from Indonesia were supplied by the Instituto The main steps of the flowering process, that is,
Nacional de Pesquisas da Amazônia (INPA), Amazon, induction, evocation, initiation and flower development
Brazil. may be differently affected by environmental factors
(Bernier et al., 1985; Kinet et al., 1985). Daylength and
Growth conditions - Seeds were germinated in a
temperature are factors known to influence this
roll of filter paper moistened with distilled water. After
process (Vince-Prue, 1975; Cockshull & Kofranek,
germination, the seedlings were transferred to pots
1994; Ramin & Atherton, 1994), as well as vegetative
containing 3 kg of a mixture of soil and sand (4:1), and
growth (Schwabe, 1956; Heuvelink & Bertin, 1994).
maintained in growth chambers or in a greenhouse.
The pots were watered with tap water and occasionally
with nutrient solution (Hoagland & Arnon, 1938). When
necessary, the plants were supported by bamboo
poles.
Experiments in growth cabinets - The plants were
maintained in growth cabinets (Conviron, Canada)
provided with fluorescent and incandescent lamps
(31W.m-2 at plant level), under controlled photoperiod
and temperature.
Experiments in greenhouse - The plants were kept
in the greenhouse, under natural photoperiod and,
when necessary, supplemented with artificial light
from incandescent lamps of 60W (0.05W.m-2) to
prolong the daylength.
Effect of photoperiod - Plants were maintained in
the greenhouse under 8h of natural light
supplemented with different periods of artificial light
(60W incandescent lamps), providing photoperiods of
10, 12, 14, 16 and 20h. Plants were harvested 45, 60
and 70 days after initiating treatments. During the
period of the experiments the mean temperatures
varied from 13oC (June) to 30oC (February).
Influence of different thermal regimes - Two
experiments were carried out in growth cabinets with
a 13h daylength and different combinations of
day/night temperatures (20/15, 25/20 and 30/25oC). In
a later experiment plants were kept under unfavorable
conditions for flowering and tuberization (determined
in previous experiments) until the plants reached 9 to FIGURE 1- Effect of different photoperiods on number
10 nodes on the main stem using a photoperiod of 20h of nodes (A) and on height (B) of the main stem of P.
and a day/night temperature of 30/25oC. After this tetragonolobus.
period, the plants were subjected to day/night

R.Bras.Fisiol.Veg., 8(2):105-110, 1996.

 107

FIGURE 2- Effect of different photoperiods on

flowering of P. tetragonolobus.

Winged bean plants submitted to a 20h photoperiod

were normally taller than plants maintained under
other photoperiods (10, 12, 14, and 16h daylight) as
result of an increase both in the number of nodes and
the elongation of the internodes (fig. 1A, B). The
influence of photoperiod on plant growth confirms
previous studies (Wong & Schwabe, 1979; Russel &
Stuber, 1983; Thomas & Raper, 1984). Wong &
Schwabe (1979) found a greater dry mass for plants
under photoperiods of 16h than those of 8h, when
plants were growing at 26/18oC day/night
temperatures.
In the greenhouse, with controlled photoperiods, the
experiments were conducted at ambient temperature
that varied from 13 to 30oC, over the experimental
period. The landrace of winged bean under
investigation may be considered a short-day plant,
since flowering occurred only under photoperiods
shorter than 20h. For the photoperiod of 16h there was
a delay in flower initiation in relation to the other
photoperiodic treatments: 50% after 45 days and 75% FIGURE 3- Effect of different day/night temperatures
after 70 days (fig. 2). Other authors have already on number of nodes on main stem (A), number of
stated that most of the winged bean varieties studied leaves (B) and on height of the main stem (C) of P.
are sensible to daylength (Herath & Ormrod, 1979), tetragonolobus (Experiment II).
and therefore classified as short-day plants (Khan,
1978).
short-day for tuberization. Root tuber formation on the
In several species the formation of underground winged bean plants only occurred for plants
reserve organs, like stems and tuberous roots, is maintained under photoperiods shorter than 16h,
dependent on or accelerated by exposure of the when in the greenhouse. Plants under a 10h
leaves to adequate photoperiods. Several authors photoperiod produced close to 3 tubers per plant,
have observed that the process of formation of these those under 12 and 16h photoperiods close to 2 tubers
organs may be enhanced by exposing plants to per plant and those under a daylength of 14h, about 4
short-days (Vince-Prue, 1975; Okubo et al, 1992). tubers per plant.
There are few studies on the influence of photoperiod
on root tuber formation. Alvarenga & Válio (1989) Considering the observations, two experiments
working with Pachyrhizus tuberosus, observed that were carried out (Exp.I and II - tab. 1, 2, 3; fig. 3). The
tuberization occurred only under photoperiods shorter plants remained for 63 or 96 days in growth cabinets
than 16h, and so they classified this species as
R.Bras.Fisiol.Veg., 8(2):105-110, 1996.
108

TABLE 1- Effect of different day/night temperatures TABLE 2- Effect of different day/night temperatures
on P. tetragonolobus growth, after 96 days in growth on tuberous root formation of P. tetragonolobus
cabinets (Experiment I) and of dry matter (Experiment I).
accumulation after 63 days in growth cabinets
Day/night temperature (oC)
(Experiment II).
20/15 25/20 30/25
Day/night temperature (oC)
Plants with tubers (%) 100 100 90
20/15 25/20 30/25
Number per plant 1a 3a 2a
Experiment I
Fresh mass (g) 1.41a 6.32b 1.37a
Plants length (mm) 510.4a 1216.4b 1255.0b
Dry mass (g) 0.32a 1.67b 0.30a
Number of leaves 13a 33b 31b
Length (mm) 61.3a 54.5a 57.5a
Number of nodes 13a 16b 15b
Bigger diameter (mm) 4.0a 8.8b 4.6a
Number of branches 1a 4b 4b
Leaf dry mass (g) 0.38a 1.58b 2.49c The values followed by different letters are significantly
different at 5% probability level, according to Tukey’s
Shoot dry mass (g) 0.62a 2.59b 5.09c test.
2
Leaf area (m ) 0.0124a 0.0685b 0.1494c
occurred with plants under a 25/20 oC day/night
Experiment II temperature. In the first experiment, flower initiation of
Shoot dry mass (g) 0.54 1.79 4.11 plants submitted to a 25/20oC day/night temperature
occurred after 48 days. After 52 days, all the plants had
The values followed by different letters are significantly flowered, but, after 89 days, all the flowers aborted. In
different at 5% probability level, according to Tukey’s the second experiment, flower initiation started after
test. 49 days and, after 55 days, all the plants were in
flower. In the first experiment, the first flower appeared
under a 13h photoperiod and different day/night between the 9th and 14th node (from the plant base).
temperature regimes (20/15, 25/20 and 30/25 oC). The In the second experiment, the first node to show an
plant growth pattern was strongly affected by inflorescence was the 11th. Plants under 20/15 and
temperature. Plants under 20/15oC day/night 30/25oC day/night temperature did not produce
temperatures exhibited a reduction in the number of flowers until harvest (96 and 63 days, respectively).
leaves, branches and nodes on the main stem. Such
plants also showed a smaller main stem length, a Temperature and photoperiod interact in such a
reduction in total dry mass and lower leaf area than manner that the critical photoperiod tends to change
plants under 25/20 and 30/25oC temperature regimes. with the temperature, as has been reported by Heide
There was no difference between the plants growing (1977) for strawberry. Wong & Schwabe (1979) found
under 25/20 and 30/25oC day/night temperatures, with that day/night temperature is as important as
regard to plant height, number of nodes and main stem daylength in controlling the flowering of winged bean.
ramifications (tab. 1 and fig. 3A, C). Under these two They reported that under the 8h daylength, plants
temperature regimes, the number of leaves was also flowered at 26/18oC whereas at a higher temperature
similar in the first experiment (tab. 1), but it was higher of 32/22oC, flowering was inhibited. In a second
under 30/25oC, in the second experiment (fig. 3B). In experiment they showed that this inhibitory effect
this experiment, plants under 30/25oC showed higher could be caused by the high day temperature, rather
dry mass accumulation and leaf area than those under than by high night temperature. Herath & Ormrod
other thermal regimes: shoot dry mass 2 times higher (1979) working with 20 different varieties of winged
and leaf 1.5 times higher than plants that had grown at bean, also observed that temperature was as
25/20oC. Plants submitted to a 30/25oC day/night important as photoperiod in controlling flowering of
temperature had, at the end of the experiment I, a leaf these plants. Seventeen of the varieties flowered
area twice that of plants grown at 25/20oC and 12 under 11h photoperiod at 25/20oC, but failed to flower
times that of those grown at 20/15oC (tab. 1). at the same photoperiod at 30/25oC and none of the
varieties flowered under a 14h photoperiod. Rueegg
Wong & Schwabe (1979) found that for a 32/22oC (1981) studied four groups of winged bean under 12h
day/night temperature there was no difference in dry photoperiod and day/night temperature of 27/23 and
mass accumulation between plants that had grown 22/18oC. Plants of all the treatments flowered and the
under 16 or 8h photoperiods. highest seed production was at 27/23 oC.
Although plants were under an inductive
photoperiod during both experiments, flowering only
R.Bras.Fisiol.Veg., 8(2):105-110, 1996.
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TABLE 3- Effect of different day/night temperatures Tuberization in Pachyrhizus tuberosus was more
on tuberous root formation of P. tetragonolobus affected by temperature than photoperiod. Higher
(Experiment II). temperatures (30/25oC) inhibited tuber formation, and
under lower temperatures (20/15 and 25/20 oC) there
Day/night temperature (oC) was an increase in the number of roots (Alvarenga &
20/15 25/20 30/25 Válio, 1989).
Plants with tubers (%) 20a 90b 0 Another experiment was carryed out (Exp. III), to
standardize the initial growth of the main stem. The
Number per plant 1a 1a -
plants were mantained under 30/25oC day/night
Dry mass (g) 0.09a 0.39b - temperatures and a 20h photoperiod until they reach
Length (mm) 67.0a 66.6a -
9 to 10 nodes on the main stem. These conditions
were, according to previous observations,
Bigger diameter (mm) 5.5a 5.3a - unfavourable for flowering and tuberization but
favorable for stem growth. After 20 days under these
The values followed by different letters are significantly
different at 5% probability level, according to Tukey’s
conditions the plants were measured (tab.4 - numbers
test. in brackets) and transferred to the day/night
temperatures previously tested. After 80 days they
were harvested and measured. Plants under the lower
TABLE 4- Height, number of nodes on main stem and
thermal regime (20/15oC) showed, again, lower stem
dry mass of shoot of plants of P. tetragonolobus that,
growth, lower number of nodes on the main stem, and
after remain 20 days under 30/25o C and 20h
lower shoot dry mass than the others. Plants under
photoperiod, were maintained for 80 days under
other thermal regimes did not show any difference for
different day/night temperatures and 13h photoperiod
the characteristics analyzed (tab. 4).
(Experiment III).
The seeds of winged bean used to carry on this work
Temperature Height of the No of nodes on Dry mass of
(oC) main stem (m) main stem the shoot (g) were obtained from Indonesia plants. This country is
localized in the equatorial region (between 11 oS and
20/15 1.108a (0,652) 15a (10) 1.70a
5o55N latitude), where there is almost no fluctuation in
25/20 2.314b (0,638) 30b (10) 8.36b photoperiod. The abundance of mountains on this
30/25 2.523b (0,591) 33b (9) 6.31b large extension of land also contributes for reduction
of annual temperatures, normally higher than 25oC, to
The means in brackets correspond to the initial height around 18oC. In our experimental conditions, the
and initial number of nodes on main stem (plants winged bean plants only produced flowers and
under 30/25 o C and 20h photoperiod for 20 days). The tuberous roots under photoperiods lower than 16
means were not significantly different. hours, and presented an increase on tuberous root dry
mass under thermal regimes of 25/20oC. These
Wong & Schwabe (1979) found that short days are characteristics coincide with those of the native habitat
necessary for winged bean to produce tubers almost of P. tetragonolobus. Thus, flowering and tuberization
irrespective of the day/night temperature. However, in this species can be controlled by photo- and
Rueegg (1981) reported that only two of the four thermoperiod.
groups studied produced tubers under short days, and
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