See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/355874112

Chlorophyll Fluorescence and Growth Evaluation of Ornamental Foliage

Plants in Response to Light Intensity Levels under Continuous Lighting
Conditions

Article  in  Flower Research Journal · November 2021

DOI: 10.11623/frj.2021.29.3.05

CITATIONS READS

0 18

4 authors, including:

Raisa Aone Marciales Cabahug

Sahmyook University
21 PUBLICATIONS   23 CITATIONS   

SEE PROFILE

Some of the authors of this publication are also working on these related projects:

Succulent and Cactus Business Project View project

Succulent Mutation View project

All content following this page was uploaded by Raisa Aone Marciales Cabahug on 07 January 2022.

The user has requested enhancement of the downloaded file.

Flower Res. J. (2021) 29(3) : 153-164 ISSN 1225-5009(Print)
DOI https://doi.org/10.11623/frj.2021.29.3.05 ISSN 2287-772X(Online)

O R IG IN A L A R T IC L E

Chlorophyll Fluorescence and Growth Evaluation of Ornamental

Foliage Plants in Response to Light Intensity Levels under Continuous
Lighting Conditions
Jae Hwan Lee1,2†, Raisa Aone M. Cabahug3†, Na Hyun You1, and Sang Yong Nam1,2*
1
Department of Environmental Horticulture, Sahmyook University, Seoul 01795, Korea
2
Natural Science Research Institute, Sahmyook University, Seoul 01795, Korea
3
Plant Genetics and Breeding Institute, Sahmyook University, Seoul 01795, Korea
†
These authors contributed equally to this work.

Received 5 September 2021; Revised 14 September 2021; Accepted 15 September 2021

Copyright © 2021 by The Korean Society for Floricultural Science

Abstract Foliage plants are considered as an integral part Introduction

of adding aesthetic and functional purposes in public places.
Nowadays, various foliage plants are used to decorate and
enhance the aesthetics of several government and private
establishments which offer and run services non-stop that
involve medical/police emergencies, entertainment, and travel.
Under these conditions, indoor foliage plants are subjected
to continuous lighting conditions. Thus, this study aimed
to determine the growth and physiological response
of common foliage plants (Hoya carnosa f. variegata,
Epipremnum aureum f. variegata, Rhapis excelsa, Hedera
helix, Chamaedorea elegans, and Spathiphyllum wallisii)
under continuous light conditions with varying light intensity
levels (60, 120, and 180 µmol m-2 s-1). Plant responses were
evaluated using growth parameters and the chlorophyll
fluorescence analysis of the OJIP curve and its specific
energy flux parameters. Results showed that foliage plants
showed positive growth and increase in mass yield in higher
light intensities and had a minor impact on color quality.
Chlorophyll fluorescence analysis suggested that strong
stress responses were evident in low light conditions,
whereas in fluorescence parameters, continuous lighting
conditions with high light intensities showed stress due to
excess light to shade-tolerant plants.
Additional key words: foliage plants, light stress, OJIP,
shade-tolerant
*Corresponding author: Sang Yong Nam
Tel: +82-2-3399-1732
E-mail: namsy@syu.ac.kr
ORCID: https://orcid.org/0000-0002-4351-447X
As one of the essential components in ornamental crops,
foliage plants are grown for their attractive leaves with
various shapes, colors, styles, and textures, which are
typically used for outdoor landscapes or interior scaping
(Henny and Chen 2003), add aesthetic beauty to a living
space (Kochlar 2009), provide functional features such as a
partitioning and screen device (Randani et al. 2017), and has
been found to improve task performance and act as a mood
booster (Shibata and Suzuki 2002). Thus, it is not a surprise
that foliage plants have continued to boost their market share
in the floriculture industry and have continued to surge in
market demands for the last decade (USDA 2019).
Aside from homes and offices, a large market of
ornamental foliage plant species is being purchased to be
cultivated and grown indoors under continuous lighting
within commercial spaces, buildings, and other public
infrastructures that operate day and night, such as malls,
hospitals, and airports. These ornamental plants have often
been placed or landscaped around attractions or relaxing
places for guests arriving at these facilities as green walls,
center islands, and plant boxes. Nowadays, smart green
buildings and homes are continuously developing and
improving technologies in assisting people and businesses
to provide favorable conditions to facilitate the growth and

www.ijfs.org
154
development of indoor plants, thus it is important to
optimize and determine plant species’ responses in these
given situations (Chen et al. 2017; Shagol et al. 2018).
One of the most challenging aspects for indoor management
for these foliage plants is the light conditions. Light is the
most critical factor influencing foliage plant performances
under interior conditions (Conover and Poole 2011), more
so for extended light periods under light-emitting diodes
(LED), as it affects both photosynthetic rate and assimilate
accumulation. Several research studies conducted on indoor
light intensities of foliage plants have ranged from < 20 -
-2 -1
300 µmol m s (Park et al. 2010a; Park et al. 2010b;
Shagol et al. 2018) which could be classified into different
categories as low, medium, and high (Pennisi 2017);
however foliage plants generally favor low to medium light
intensities.
Depending on plant tolerance, plant responses to continuous
lighting may cause varying effects such as foliar chlorosis,
limited or reduced plant growth and productivity, sudden
onset of senescence, and early flowering (Sysoeva et al. 2010;
Van Gestel et al. 2005). On the other hand, other studies
have shown that continuous light exposure instigated a higher
and faster developmental rate in spring wheat (Zhukov and
Romanovskaja 1980), chickpeas (Sethi et al. 1981), and
Arabidopsis (Massa et al. 2007). However, reviews of Sysoeva
et al. (2010) suggest that light intensity is another modifying
factor that affects continuous lighting regimes which must also
be taken into consideration.
Mechanisms fundamental to the tolerance of interior light
intensities vary among different types of foliage plants
(Chen et al. 2004). Low light intensities were generally
reported to increase leaf drop and reduce leaf quality
(Conover and Poole 1997; Sawwan and Ghunem 1999),
however, other studies suggest otherwise and have increased
fresh and dry weights (Shen and Seeley, 1983) and higher
chlorophyll content (Kubatsch et al. 2007; Lance and Guy
1992). Thus, proper trials must be done on foliage types
and species in order to determine their response and
appropriate light intensities that would allow them to thrive
www.ijfs.org
Flower Res. J. (2021) 29(3) : 153-164
in indoor conditions.
An important tool that is widely used and powerful in
both basic and applied research among plant physiological
studies is the use of chlorophyll fluorescence analysis
(Kupper et al. 2019). This analysis is based on the principle
that light energy absorbed by chlorophyll molecules may be
used for photosynthesis (photochemistry), dissipated as heat
(non-photochemical quenching), or re-emitted as light
(chlorophyll fluorescence), and by determining the yield of
the latter data, information about the efficiency of the other
metabolic processes can be attained (Maxwell and Johnson
2000). This data is done to determine the effects of
physiological stresses in various economic plants such as
Aloe vera (Hazrati et al. 2016), roses (Bayat et al. 2020),
lotus (Bordenave 2019), fern (Oliwa and Skoczowski 2019),
etc.
Although light quality and duration have been often
studied among ornamental crops, continuous lighting and
light intensity levels have had scant attention despite the
increasing exposure of foliage plants to such indoor abiotic
conditions. Hence, this study deemed to determine the
impact on plants’ performance, chlorophyll fluorescence,
and growth under continuous lighting and different light
intensity levels in various foliage plants.
Materials and Methods
Plant materials
Six ornamental foliage plants, namely Hoya carnosa f.
variegata, Epipremnum aureum f. variegata, Rhapis excelsa,
Hedera helix, Chamaedorea elegans, and Spathiphyllum
wallisii (Fig. 1) which were taken from a plant nursery in
Gyeonggi-do, South Korea. Experimental units were then
taken to the Plant Physiology Laboratory, Department of
Environmental Horticulture, Sahmyook University.
Before the conduct of the experiment, plants were
repotted into premixed horticultural soil (Shinsung Mineral,
South Korea) to homogenize planting media. These were
then placed in the greenhouse for 3 weeks to allow the
establishment of plants into the new media.
Flower Res. J. (2021) 29(3) : 153-164 155

Fig. 1. Foliage plants subjected to continuous lighting conditions at different light intensities: A) Hoya carnosa f. variegata; B) Epipremnum
aureum f. variegata; C) Rhapis excelsa; D) Hedera helix; E) Chamaedorea elegans; and F) Spathiphyllum wallisii.

Experimental design and treatment values indicating the lightness, hue, and saturation, respectively.
The study was conducted in Completely Randomized In addition, CIELAB values were processed using the CIELAB–
Design (CRD) with three light intensity levels of 60, 120, RHS converter (CIELAB-RHS Colour Converter, Oregon State
-2 -1 University, USA) in order to determine the Royal Horticulture
and 180 µmol m s under continuous lighting conditions
using white LED bars (CPH40W, Pulse Tech, South Korea) Society (RHS) color value and group.
with 6500 K color temperature which was installed in a
SPAD chlorophyll content measurement
growth facility. These treatments were replicated thrice with
Chlorophyll content was determined using a portable SPAD
three plants per replication.
chlorophyll meter (SPAD-502, Konica Minolta, Japan). Calibration
Potted foliage plants were placed in different growing
was done by carefully clamping the instrument onto the
facilities with their respective light intensity treatments.
central part of the leaf to obtain chlorophyll readings. In each
Conditions were set at 20/18°C (± 1°C) D/N with a relative
plant, three leaves were tagged as representative and served
humidity of 75% (± 10%) for 4 weeks from March to April
as replications. SPAD readings were taken in a two-week
2021.
interval on the same leaves.
Plant growth parameters
Chlorophyll fluorescence measurement
Growth and quality evaluation was done by collecting plant
Using a portable fluorometer (FluorPen FP 110/D, Photon
height, width, fresh and dry weight. For determining the dry
Systems Instruments, Czech Republic), fluorescence transient
weight of samples, leaves and roots were oven-dried at 85°C
data were taken from fully developed leaves which were
for 24-hours wherein the weight became constant. To
selected and tagged for measurement. The leaves were
determine ornamental quality, leaf color was assessed using
dark-adapted for 30 min before starting the measurements
the CIELAB color value using a spectrophotometer (CM-2600d,
as a standard protocol indicated by the manufacturer.
Konica Minolta, Japan) which utilizes L*, a*, and b* color space

www.ijfs.org
156 Flower Res. J. (2021) 29(3) : 153-164

Table 1. Parameters, formula, and/or description of the steps for fluorescence induction (Modified from Kasemsap 2014; Stirbet and
Govindjee 2011).
Parameter Formula Description
Fluorescence transient O-J-I-P
FO Fluorescence intensity at O-step (50 μs) of O-J-I-P
Fj FJ = F2ms Fluorescence intensity at J-step (2 ms) of O-J-I-P
Fi FI = F60ms Fluorescence intensity at I-step (60 ms) of O-J-I-P
FP (= FM) Maximal fluorescence intensity, at the peak P of O-J-I-P
FV FV = FM – F0 (Maximal variable fluorescence)
Vj Vj = (Fj – F0)/(FM - F0) Relative variable fluorescence at the J-step
Vi Vi = (Fi – F0)/(FM - F0) Relative variable fluorescence at the I-step

Specific energy fluxes (per QA-reducing P II reaction center - RC)

ABS/RC ABS/RC = MO(1/VJ)(1/φPo) Absorption flux per RC
TRO/RC TRO/RC = MO(1/VJ) Trapped energy flux per RC (at t = 0)
ETO/RC ETO/RC = (MO/VJ)(1-VJ) Electron transport flux from QA to QB per RC (at t = 0)
DIO/RC DIO/RC = (ABS/RC)-(TRO/RC) Dissipated energy flux per RC (at t = 0)

Measurements were done three times for each replication on
the adaxial leaf surface at 13:00 o’clock in the afternoon at
a 2-week interval.
Parameters of the OJIP fluorescence, indicating transient
and specific energy fluxes values, were calculated as shown
in Table 1 with their respective formulas and further
descriptions. These measurements were adapted from the
studies of Stirbet and Govindjee (2011) and Kasemsap (2014).
Data analysis
Data from plant growth parameters, chlorophyll content,
and chlorophyll fluorescence measurement were organized
and subjected to standard analysis of variance (ANOVA) using
SPSS Statistics 22.0 (IBM, USA). To compare mean differences,
Duncan’s multiple range test was applied with a 5% level of
significance. Likewise, the comparison of photon yield and
specific energy fluxes were presented in radar charts using
Microsoft Excel (Microsoft, USA), while OJIP fluorescence
transient peaks were derived from the FlourPen software
(Photon Systems Instruments, Czech Republic).
Results and Discussion
Plant growth and CIELAB values
Different levels of light intensity under continuous lighting
conditions significantly affected the respective growth parameters
of various foliage plants (Table 2).
Hoya carnosa f. variegata had significantly the tallest
(18.10 cm) and bigger plants (27.24 cm) when subjected to
180 µmol m-2 s-1 and were also found to have the highest
values in terms of their dry weight (shoot and root) and
shoot fresh weight. These were then followed by plants
grown with 120 and 60 µmol m-2 s-1.
Growth parameters collected from Epipremnum aureum
f. variegata were significantly affected by lighting conditions.
Plants grown under 180 µmol m-2 s-1 had the tallest plants
(27.54 cm), fresh weight of shoots (63.34 g) and roots
(27.89 g), and dry weight of shoots (7.11 g) and roots
(3.38 g).
The highest values of growth parameters were observed
from Hedera helix from those of the 180 µmol m-2 s-1 group.
Treatments at higher light intensities (180 and 120 µmol m-2 s-1)
had significantly the tallest plants. The highest shoot fresh
and dry weight were taken from those treated at 180
µmol m-2 s-1 which did not significantly differ from those
at 120 µmol m-2 s-1, followed by those of 60 µmol m-2 s-1.
Significant differences were found in the root dry weight
with the highest value (1.00 g) taken at 180 µmol m-2 s-1
which significantly differed from those of 120 (0.81 g) and

www.ijfs.org
Flower Res. J. (2021) 29(3) : 153-164 157

Table 2. Effects of continuous lighting condition at different lighting intensities on the plant parameters, fresh and dry weight of various
foliage plants.
Plant parameters Fresh weight Dry weight
Light intensity
Species (cm) (g) (g)
(µmol m-2 s-1)
Height Width Shoot Root Shoot Root
z
Hoya carnosa f. variegata 60 16.81 c 25.07 c 78.67 c 5.25 6.09 b 1.05 c
120 17.19 b 26.10 b 81.75 b 5.96 6.67 b 1.18 b
180 18.10 a 27.24 a 91.48 a 5.79 7.46 a 1.30 a
y
F-test * * * NS * *

Epipremnum aureum f. variegata 60 26.99 b 27.10 b 56.36 b 21.99 b 5.79 c 2.61 c

120 25.60 c 30.32 a 58.61 b 21.38 b 6.92 b 2.90 b
180 27.54 a 23.40 c 63.34 a 27.89 a 7.11 a 3.38 a
F-test * * * * * *

Rhapis excelsa 60 36.56 c 28.09 14.59 8.54 6.25 4.38

120 38.66 a 29.79 15.44 8.69 6.60 4.39
180 37.86 b 30.21 14.96 6.57 6.42 3.49
F-test * NS NS NS NS NS

Hedera helix 60 24.46 c 29.31 18.18 b 2.52 4.60 b 0.71 b

120 26.79 b 32.63 23.79 a 2.59 6.14 a 0.81 b
180 28.10 a 27.26 25.50 a 3.11 6.94 a 1.00 a
F-test * NS ** NS ** *

Chamaedorea elegans 60 27.40 b 23.77 a 11.33 b 3.88 b 3.53 1.54

120 28.16 a 22.97 b 14.06 a 4.55 a 4.22 1.72
180 27.24 b 19.79 c 12.12 b 3.62 b 3.80 1.52
F-test * ** * * NS NS

Spathiphyllum wallisii 60 36.49 b 27.81 b 40.55 c 14.23 6.75 b 2.37 c

120 35.87 b 24.03 c 42.37 b 12.87 6.81 b 2.94 b
180 40.16 a 28.63 a 43.34 a 13.12 7.43 a 3.16 a
F-test * * * NS * *
z
Mean separation within columns between treatments by Duncan’s multiple range test at p = 0.05.
y
NS, *, **; Non-significant, or significant at p = 0.05 or 0.01, respectively.

60 (0.71 g) µmol m-2 s-1. Likewise, similar trends were
observed from those of Spathiphyllum wallisii which
statistically had the tallest (40.16 cm) and bigger plants
(28.63 cm), and the highest value for shoot fresh weight
(43.34 g), and shoot (7.43 g) and root (3.16 g) dry weight.
Rhapis excelsa foliage plants had significantly the tallest
plants treated under 120 µmol m-2 s-1 with 38.66 cm, followed
-2 -1
by those grown under 180 and 60 µmol m s with 37.86
cm and 36.56 cm, respectively. These were similar to those
of their fresh and dry weights, however, mean differences
were found to be insignificant. Likewise, Chamaedorea
elegans also had favorable growth under 120 µmol m-2 s-1 light
intensity which had the tallest plants (28.16 cm), and highest
shoot (14.06 g) and root (4.55 g) fresh weight. These were
followed by plants grown under 60 and 180 µmol m-2 s-1,
which did not significantly differ from each other.
Although foliage plants are generally tolerant to shaded
conditions, this ability of plants can be traced in their

www.ijfs.org
158 Flower Res. J. (2021) 29(3) : 153-164

efficiency to utilize light energy during photosynthesis and
the rate of respiration under different low light intensities
(Corre 1983). Under continuous lighting conditions, the use
of a 24-h photoperiod with relatively low photosynthetic
photon flux (PPF) has been reported to have benefited
tolerant crops (Sysoeva et al. 2010). The increase of plant
growth including fresh and dry weight favoring higher light
intensities is indicative of the treatment’s importance to the
physiological processes involved in its development (Chen
et al. 2004). Taiz and Zeiger (1991) explain that at higher
lux/PPF values, the movement of the chloroplast is parallel
to the indecent light increasing plant height. In addition, it
has been well established that appropriate high light
exposures would enhance photosynthetic activity which
would, in turn, have higher growth rates (Adams and Early
2004). Similar studies have been observed in other potted
ornamental plants such as in succulents (Cabahug et al.
2017; Nam et al. 2016) and H. helix (Park et al. 2010a).

Table 3. CIELAB values indicating the RHS Color group and value as affected by continuous lighting conditions at different lighting
intensities of various foliage plants.
Light intensity CIELAB color values
Cultivar RHS color group RHS value
(µmol m-2 s-1) L* a* b*
Hoya carnosa f. variegata 60 32.57 ± 0.66 -5.76 ± 0.37 9.40 ± 0.84 Yellow-Green 147A
120 33.77 ± 0.59 -5.88 ± 0.27 10.19 ± 0.66 Yellow-Green 147A
180 34.67 ± 0.73 -5.49 ± 0.34 11.08 ± 0.66 Yellow-Green 147A
y
F-test NS NS NS

Epipremnum aureum f. variegata 60 32.89 ± 0.51 bz -8.01 ± 0.25 a 12.36 ± 0.67 c Green N137A
120 36.13 ± 0.70 a -9.24 ± 0.17 b 15.87 ± 0.74 b Green N137B
180 36.80 ± 0.50 a -8.14 ± 0.17 a 16.48 ± 0.72 a Green N137B
F-test ** ** **

Rhapis excelsa 60 32.24 ± 0.56 b -6.89 ± 0.25 b 13.09 ± 0.80 b Yellow-Green 147A
120 31.22 ± 0.63 b -5.90 ± 0.22 a 11.35 ± 0.63 b Yellow-Green 147A
180 34.51 ± 0.93 a -6.35 ± 0.16 b 16.02 ± 1.19 a Green N137A
F-test ** ** **

Hedera helix 60 37.77 ± 0.77 -9.25 ± 0.35 16.60 ± 1.12 Green N137B
120 38.68 ± 0.96 -9.11 ± 0.43 18.08 ± 1.41 Green N137D
180 38.41 ± 1.01 -9.24 ± 0.34 18.44 ± 1.10 Green N137D
F-test NS NS NS

Chamaedorea elegans 60 35.99 ± 0.73 -8.52 ± 0.19 19.12 ± 0.73 Green N137B
120 35.67 ± 0.83 -8.34 ± 0.29 19.64 ± 1.05 Green N137B
180 36.03 ± 0.70 -7.99 ± 0.22 19.93 ± 0.71 Green N137B
F-test NS NS NS

Spathiphyllum wallisii 60 35.99 ± 0.73 -8.52 ± 0.19 19.12 ± 0.73 Green N137B
120 35.67 ± 0.83 -8.34 ± 0.29 19.64 ± 1.05 Green N137B
180 36.03 ± 0.70 -7.99 ± 0.22 19.93 ± 0.71 Green N137B
F-test NS NS NS
z
Mean separation within columns between treatments by Duncan’s multiple range test at p = 0.05.
y
NS, *, **; Non-significant, or significant at p = 0.05 or 0.01, respectively.

www.ijfs.org
Flower Res. J. (2021) 29(3) : 153-164 159

However, in more shade-tolerant plants, wherein their light
saturation point is lower, even at low light levels would
prompt an increase in height and leaf expansion as means
to adapt to low light conditions (Steinger et al. 2003; Zhang
et al. 2003).
Based on the results of the CIELAB values, significant
color differences were observed from E. aureum f. variegata
and R. excelsa. Results showed that brighter, lighter hue,
and more saturated in color at higher light intensities.
Among foliage plants, only those of E. aureum f. variegata
Lower light intensities were tagged as yellow-green while at
the highest level was categorized as green.
Changes in colors for ornamental foliage plants have
been of great importance due to their aesthetic value.
Studies by Kim et al. (2012) suggested that some foliage
plants under low light conditions may have less or no leaf
color changes at all.
Chlorophyll content
Chlorophyll content (SPAD units) was significantly affected

and R. excelsa have been significantly affected by different by the different light intensities under continuous lighting

lighting intensities in their CIELAB values as shown in Table conditions as shown in Fig. 2.

3. E. aureum f. variegata plants which were treated under Results showed that most of the foliage plants, H.

180 µmol m-2 s-1 significantly had the highest a* and b* carnosa f. variegata, H. helix, C. elegans, and S. wallisii

values with -8.14 and 16.48, respectively. In their L* values, had significantly higher chlorophyll content when subjected

those grown at the highest light intensities had significantly to 180 µmol m-2 s-1, which was then followed by those treated

differed from other light levels with 36.80. However, based at 60 and 120 µmol m-2 s-1. On the other hand, E. aureum f.

on the RHS color group and value, they are still categorized variegata had the highest chlorophyll content from those of

as green. R. excelsa plants, on the other hand, had 60 µmol m-2 s-1 (58.21), followed by those of 120 and 180

significantly changed in RHS color group and value which µmol m-2 s-1 with 52.76 and 49.45 which did not statistically

is indicative of its high CIELAB values at 180 µmol m-2 s-1. differ from each other. R. excelsa plants grown under 120

Fig. 2. Chlorophyll content (SPAD units) of selected foliage plants grown under continuous lighting conditions at different light intensities.
Error bars in each column show standard error values. Mean separation within columns between treatments by Duncan’s multiple range
test at p = 0.05. NS, *, **; Non-significant, or significant at p = 0.05 or 0.01, respectively.

www.ijfs.org
160 Flower Res. J. (2021) 29(3) : 153-164

µmol m-2 s-1 significantly had the highest chlorophyll content
of 61.40, followed by those subjected at 60 and 180 µmol
-2 -1
m s with 57.16 and 54.70, respectively.
One of the most essential factors wherein light condition
affects plants is the leaf pigment. Color-leaf plants are found
to be sensitive to light and mostly produce anthocyanins in
response to the light being absorbed especially as a response
to environmental changes (Chen et al. 2008; Michal 2009).
According to Zhao et al. (2012), low-light environments are
likely to interfere with normal photosynthetic processes by
affecting the synthesis and ratio of various plant pigments
which include chlorophyll. Research studies dealing with low
light intensity largely reported that as light intensity decreases
continuous lighting conditions. Langton et al. (2003) suggested
that, in some plant species, prolonged photoperiod also
results in increases in leaf area and chlorophyll content per
unit of leaf area which would mirror the increase in biomass
production. This trend in continuous light conditions has
been observed in the production of mini-cucumbers (Lanoue
et al. 2021), Platycodon grandiflorum (Xiao et al. 2007),
among others. Notably, reviews of Sysoeva et al. (2010) reported
that leaf pigment content as affected by photoperiod has
been found to be varied depending on species.
Chlorophyll fluorescence analysis
According to the reviews of Stirbet and Govindjee (2011),

chlorophyll contents significantly increases such as that of OJIP chlorophyll transient has been able to determine and

Physocarpus species (Zhang et al. 2016), Brassica campestris monitor the effects of environmental stressors on the

(Zhu et al. 2017), Sage (Rezai et al. 2018), and other foliage photosynthetic apparatus and their respective functions.

plants such as Dieffenbachia, Anthurium, and Ficus (Chen et Each step of the OJIP curve corresponds to specific

al. 2005), which was attributed to the correlated increase of activities involved in the photosystems. Starting from the

specific leaf area and enlarging leaf area. initial fluorescence (Fo) or the level O, the induction curve

However, this has not been the case for plants under appears toward levels J and I emitted after 2 and 60 ms,

Fig. 3. Comparison of OJIP fluorescence transient of various foliage plants: A) Hoya carnosa f. variegata; B) Epipremnum aureum f.
variegata; C) Rhapis excelsa; D) Hedera helix; E) Chamaedorea elegans; and F) Spathiphyllum wallisii exposed to continuous lighting
under different light intensities.

www.ijfs.org
Flower Res. J. (2021) 29(3) : 153-164 161

which is then followed by Fm or level P (Strasser and
Strasser 1995). The reduction of the acceptors of PS II is
indicated by the O-J section, reduction of plastoquinone
(PQ) pool by the J-I section, and reduction of acceptors of
PS I by the I-P section (Ripoll et al. 2016).
Results on the effect of different light intensities under
continuous lighting conditions are shown in Fig. 3. The
relative fluorescence intensity in the leaves of plants treated
-2 -1
under the lowest light intensity (60 µmol m s ) was higher
-2 -1
than that of 120 and 180 µmol m s , respectively, from all
foliage species at all points in the OJIP curve. Moreover,
evident differences from light intensity levels are observed
starting from the J, I, and P peaks. The increased values of
the fluorescence value at these points, especially at points
J and I, indicate a reduced photochemical activity of the PSII
(Zhang et al. 2016). For R. excelsa (Fig. 3c), C. elegans (Fig.
3e), and S. wallisii (Fig. 3f), the large decrease in the
-2 -1
fluorescence value of 120 and 180 µmol m s resulting in
a flatter OJIP curve. This was, likewise, observed among H.
-2 -1
helix at 180 µmol m s (Fig. 3b).
Differences in the radial plot analysis were observed from
OJIP parameters and energy fluxes were mostly observed
from those of Fm/Fo, Fv/Fo, ABS/RC, and DIO/RC as affected
by light intensity levels under continuous lighting conditions
(Fig. 4). The Fm/Fo and Fv/Fo were higher at lower light
intensities. These parameters are indicative of chlorophyll
fluorescence quenching activities which are often associated
with stress-induced changes in photosynthetic activities (Cen
et al. 2017). Dan et al. (2000) suggested that Fv/Fo is a good
indicator of the number and the size of photosynthetic
reaction centers. Although there was a decline of these
parameters in higher light intensities, these may have been
attributed to the continuous light exposure. Although continuous
light has been applauded for short-term exposure, the
long-term impacts on photosynthesis have been reported to
be contradictory citing a reduction in photosynthetic activity
resulting in an inefficient accumulation of starch, imbalance
of source-sink ratio, etc. (Sysoeva et al. 2010). Similar
results were found in continuous light treated mung bean
in which decline of the Fv/Fo values was attributed to the

Fig. 4. Radial plot analysis of OJIP fluorescence transient parameters and specific energy fluxes of various foliage plants: A) Hoya carnosa
f. variegata; B) Epipremnum aureum f. variegata; C) Rhapis excelsa; D) Hedera helix; E) Chamaedorea elegans; and F) Spathiphyllum
wallisii exposed to continuous lighting under different light intensities.

www.ijfs.org
162
shift in the rate of electron transport from PSII to the
principal electron acceptors with the decrease in the
number and size of the RC (Kumar et al. 2020).
Specific energy fluxes (i.e., ABS, TR, and ET) are expressions
of fully active PS II reaction center (RC). Among specific energy
flux parameters, the absorption flux per RC (ABS/RC) and
dissipated energy flux per RC (DIo/RC) had higher values in
180 µmol m-2 s-1 compared to those grown under lower
light conditions (60 and 120 µmol m-2 s-1). ABS/RC is said
to be the measure of the average absorption per active RC
and of the average amount of absorbing chlorophylls per
RC, i.e., of the apparent antenna size (Strasser et al. 2004).
On the other hand, DIo/RC denotes the proportion of the
total dissipation of untrapped excitation energy from all active
RCs, wherein the dissipation transpires as heat, fluorescence,
and energy transfer to other systems (Grieco et al. 2015).
Opposing views in some studies dealing with the values
of ABS/RC and DIo/RC. Studies by Zhang et al. (2016)
resulted in an increase in its value as an adaptive response
to low light conditions which have been attributed to the
enhanced activity of antenna pigments per reaction center.
Results of the studies of Kumar et al. (2020) indicated that
under continuous light conditions there would be a decrease
value indicating an increased number of active centers.
Likewise, similar explanations are both given for the values
of DIo/RC. Given that the foliage plants used in this study
are considered to be low-light tolerant, the increase of
ABS/RC and DIo/RC is attributed to the exposure to excess
irradiance causing the inactivity of reaction centers.
In summary, foliage plants showed a stronger response to
continuous lighting conditions at higher light intensities. Plant
parameters were found to have favorable growth and mass
yield in higher light intensities. However, differences in light
intensities under continuous light conditions did not affect the
hue, saturation, and lightness of colors of the majority of
foliage plants. In evaluating the OJIP curve, suggest strong
stress responses were evident in low light conditions. However,
in the analysis of fluorescence parameters, continuous lighting
conditions with high light intensities may provide stress to
www.ijfs.org
Flower Res. J. (2021) 29(3) : 153-164
low-light tolerant plants. A future prospective study may be
deemed necessary for investigating other ornamental plants
to determine responses in such conditions.
Acknowledgement
This research was supported by the Sahmyook University
Research Fund and the Korean Institute of Civil Engineering
and Building Technology.
References
Adams CR, Early MP (2004) Principle of horticulture. Elsevier
Butterworth-Heinemann, London, England
Bayat L, Arab M, Aliniaeifard SA, Seif M, Lastochkina O, Li
T (2018) Effects of growth under different light spectra on
the subsequent high light tolerance in rose plants. AoB
Plants 10:1-17 DOI:10.1093/aobpla/ply052
Bordenave CD, Rocco R, Maiale SJ, Campestre MP, Ruiz OA,
Rodriguez AA, Menedez AB (2019) Chlorophyll a
fluorescence analysis reveals divergent photosystem II
responses to saline, alkaline and saline-alkaline stresses
in the two Lotus japonicus model ecotypes MG20 and
Gifu-129. Acta Physiol Plant 41:167 DOI:10.1007/s11738-
019-2956-0
Cabahug RAM, Soh SY, Nam SY (2017) Effects of light
intensity on the growth and anthocyanin content of
Echeveria agavoides and E. marcus. Flower Res J
25:262-269 DOI:10.11623/frj.2017.25.4.11
Cen H, Weng H, Yao J, He M, Lv J, Hua S, Li H, He Y (2017)
Chlorophyll fluorescence imaging uncovers photosynthetic
fingerprint of citrus Huanglongbing. Front Plant Sci 8:1509
DOI:10.3389/fpls.2017.01509
Chen J, Wang Q Henny RJ, McConnell DB (2005) Response
of tropical foliage plants to interior low light conditions.
Acta Hort 669:51-56
Chen LS, Li P, Cheng L (2008) Effects of high temperature
coupled with high light on the balance between
photooxidation and photoprotection in the sun-exposed
peel of apple. Planta 228:745-756 DOI:10.1007/s00425-
008-0776-3
Chen M, Chory J, Fankhauser C (2004) Light signal transduction
in higher plants. Annual Review of Genetics 38:87-117
Chen M, Yang J, Zhu X, Wang X, Liu M, Song J (2017) Smart
Home 2.0: Innovative smart home system powered by
botanical IoT and emotion detection. Mob Netw Appl
22:1159-1169 DOI:10.1007/s11036-017-0866-1
Flower Res. J. (2021) 29(3) : 153-164
Conover CA, Poole RT (1977) Effects of cultural practice on
acclimatization of Ficus benjamina L. J Amer Soc Hort Sci
102:529-531
Conover CA, Poole RT (2011) Acclimatization of indoor foliage
plants. Hort Rev 119-154 DOI:10.1002/9781118060797.
ch4
Corre WJ (1983) Growth and morphogenesis of sun and
shade plants: The influence of light intensities. Acta Bot
Neerl 32:49-62
Dan TV, Krishnaraj S, Saxena PK (2000) Metal tolerance of
scented geranium (Pelargonium sp. ‘Frensham’): Effects of
cadmium and nickel on chlorophyll fluorescence kinetics.
Int J Phytoremediat 2:91-104 DOI:10.1080/1522651000
8500032
Grieco M, Suorsa M, Jajoo A, Tikkanen M, Aro EM (2015)
Light-harvesting II antenna trimers connect energetically
the entire photosynthetic machinery-including both
photosystems II and I. Biochim Biophys Acta Bioenerg
1847:607-619 DOI:10.1016/j.bbabio.2015.03.004
Hazrati S, Tahmasebi-Sarvestani Z, Modarres-Sanavy SA,
Mokhtassi-Bidgoli A, Nicola S (2016) Effects of water
stress and light intensity on chlorophyll fluorescence
parameters and pigments of Aloe vera L. Plant Physiol
Biochem 106:141-148 DOI:10.1016/j.plaphy.2016.04.046
Henny RJ, Chen JJ (2003) Cultivar development of
ornamental foliage plants. In: Janick J (ed) Plant breeding
reviews. John Wiley & Sons, New Jersey, pp 246-248
Kalaji MH, Guo P (2008) Chlorophyll fluorescence: A useful
tool in barley plant breeding programs. In: Sanchez A,
Gutierrez SJ (eds) Photochemistry research progress.
Nova Publishers, New York, pp 439-463
Kim J, Kang SW, Pak CH, Kim MS (2012) Changes in leaf
variegation and coloration of English ivy and polka dot
plant under various indoor light intensities. Hort Tech
22:49-55
Kochlar SL (2009) Economic botany in the tropics. 3rd ed,
Macmillan Publishers, Indian Ltd., New Delhi
Kubatsch A, Gruneberg H, Ulrichs C (2007) The effect of low
light intensity and temperature on growth of Schefflera
arboricola in interior landscapes. HortScience 42:65-67
DOI:10.21273/HORTSCI.42.1.65
Kumar D, Singh H, Raj S, Soni V (2020) Chlorophyll a
fluorescence kinetics of mung bean (Vigna radiata L.)
grown under artificial continuous light. Biochem Biophys
Rep 24:100813 DOI:10.1016/j.bbrep.2020.100813
Kupper H, Benedikty Z, Morina F, Andersen E, Mishra A,
Trtilek M (2019) Analysis of OJIP chlorophyll fluorescence
kinetics and qa reoxidation kinetics by direct fast imaging.
Plant Physiol 179:369-381 DOI:10.1104/pp.18.00953
www.ijfs.org
163
Lance CJ, Guy CL (1992) Chances in pigment levels, rubisco
and respiratory enzyme-activity of Ficus benjamina during
acclimation to low irradiance Physiol Plant 86:630-638
Langton FA, Adams SR, Cockshull KE (2003) Effects of
photoperiod on leaf greenness of four bedding plant
species. J Hort Sci Biotechnol 78:400-404
Lanoue J, Zheng J, Little C, Grodzinski B, Hao X (2021)
Continuous light does not compromise growth and yield
in mini-cucumber greenhouse production with supplemental
LED light. Plants 2021 10:378 DOI:10.3390/plants1002
0378
Lu CM, Jiang GM, Wang BS, Kuang TY (2003) Photosystem
II photochemistry and photosynthetic pigment composition
in salt-adapted halophyte Artimisia anethifolia grown under
outdoor conditions. J Plant Physiol 160:403-408 DOI:
10.1078/0176-1617-00839
Massa G, Eddy R, Mitchell C (2007) 101 ways to grow
Arabidopsis. Plant-Growth Facilities Website. Accessed
07 Jul. 2019, http://www.hort.purdue.edu/hort/facilities/gr
eenhouse/101exp
Maxwell K, Johnson GN (2000) Chlorophyll fluorescence-a
practical guide. J Exp Bot 51:659-668 DOI:10.1093/
jexbot/51.345.659
Michal OS (2009) Does anthocyanin degradation play a
significant role in determining pigment concentration in
plants. Plant Science 177:310-360 DOI:10.1016/j.plantsci.
2009.06.015
Nam SY, Lee HS, Soh SY, Cabahug RAM (2016) Effects
of supplementary lighting intensity and duration on
hydroponically grown Crassulaceae species. Flower Res
J 24:1-9 DOI:10.11623/frj.2016.24.1.1
Oliwa J, Skoczowski A (2019) Different response of photosynthetic
apparatus to high–light stress in sporotrophophyll and nest
leaves of Platycerium bifurcatum. Photosynthetica 57:
148-159 DOI:10.32615/ps.2019.037
Park SA, Kim MG, Yoo MH, Oh MM, Son KC (2010a) Plant
physiological responses in relation to temperature, light
intensity, and co2 concentration for the selection of
efficient foliage plants on the improvement of indoor
environment. Hort Sci Technol 28:928-936
Park SA, Kim MG, Yoo MH, Oh MM, Son KC (2010b)
Comparison of indoor CO2 removal capability of five
foliage plants by photosynthesis. Korean J Hort Sci
Technol 28:864-870
Randani MMN, Caldera HIU, Kaumal MN (2017) Screening
foliage plants for indoor air pollution abatement. Proceedings
of the 22nd International Forestry and Environment
Symposium, University of Sri Jayewardenepura, Sri Lanka
Rezai S, Etemadi N, Nikbakht A, Yousefi M, Majidi MM (2018)
164
Effect of light intensity on leaf morphology, photosynthetic
capacity, and chlorophyll content in Sage (Salvia officinalis
L.). Hort Sci Technol 36:46-57 DOI:10.12972/kjhst.
20180006
Ripoll J, Bertin N, Bidel LPR, Urban L (2016) A user’s view
of the parameters derived from the induction curves of
maximal chlorophyll a fluorescence: Perspectives for
analyzing stress. Front Plant Sci 7:1679 DOI:10.3389/
fpls.2016.01679
Sawwan JS, Ghunem RS (1999) Light acclimatization of
Schefflera arboricola Adv. Hort Sci 13:151-155
Sethi SC, Byth DE, Gowda CL, Green JM (1981) Photoperiodic
response and accelerated generation turnover in chickpea.
Field Crops Res 4:215-225
Shagol CC, Kim KJ, Han SW, Jeong NR, Kim HJ, Jung YB,
Yun HK (2018) Identification and classification of indoor
plants according to light intensity requirements for
botanical IoT application. J People Plants Environ 5:
329-341 DOI:10.11628/ksppe.2018.21.5.329
Shen GW, Seeley JG (1983) The effect of shading and nutrient
supply on variegation and nutrient content of variegated
cultivars of Peperomia obtusifolia. J Amer Soc Hort Sci
108:429-433
Shibata S, Suzuki N (2002) Effects of the foliage plant on task
performance and mood. J Environ Psych 22:265-272
DOI:10.1006/jevp.232
Steinger T, Roy BA, Stanton ML (2003) Evolution in stressful
environments II: Adaptive value and costs of plasticity in
response to low light in Sinapis arvensis. J Evol Biol
16:313-323
Stirbet A, Govindjee G (2011) On the relation between the
Kautsky effect (chlorophyll a fluorescence induction) and
Photosystem II: Basics and applications of the OJIP
fluorescence transient. J Photochem Phobiol B 104:
236-257
Strasser BJ, Strasser RJ (1995) Measuring fast fluorescence
transients to address environmental Questions: The
JIP-test. In: Mathis P (ed) Photosynthesis: From light to
biosphere. Kluwer Academic Publishers, Dordrecht,
Netherlands, pp 977-980
Strasser RJ, Tsimilli-Michael M, Srivastava A (2004) Analysis
of the chlorophyll a fluorescence transient. In: Papageorgio
GC, Govindjee G (eds) Chlorophyll a fluorescence: A
signature of photosynthesis. Springer, Netherlands, pp
www.ijfs.org
View publication stats
Flower Res. J. (2021) 29(3) : 153-164
323-362
Sysoeva MI, Markovskaya EF, Shibaeva TG (2010) Plants
under continuous light: A review. Plant Stress 4:5-17
Taiz L, Zeiger E (1991) Photosynthesis: Physiological and
ecological considerations. Plant Physiology. The Benjamin/
Cummings Publishing Company, Inc, pp 249-262
The AA, Kasemsap P (2014) Quantification of OJIP
Fluorescence Transient in Tomato plants under acute
ozone stress. Kasetsart J (Nat Sci) 48:665-675
United States Department of Agriculture(USDA) (2019)
Floriculture crops. Accessed 20 Jul. 2021, https://www.
nass.usda.gov/Publications/Todays_Reports/reports/flora
n19.pdf
Van Gestel NC, Nesbit AD, Gordon EP, Green C, Pare W,
Thompson L, Peffley EB, Tissue DT (2005) Continuous
light may induce photosynthetic downregulation in
onion-consequences for growth and biomass partitioning.
Physiol Plant 125:235-246 DOI:10.1111/j.1399-3054.2005.
00560.x
Xiao Y, Zhang Y, Dang K, Wang D (2007) Effects of
continuous 24-hour photoperiod on growth and
photosynthesis in Platycodon grandiflorum (Jacq.) A. DC.
Plants. Propag Ornam
Zhang H, Zhong H, Wang J, Sui X, Xu N (2016) Adaptive
changes in chlorophyll content and photosynthetic
features to low light in Physocarpus amurensis Maxim and
Physocarpus opulifolius “Diabolo”. PeerJ 4:e2125 DOI:
10.7717/peerj.2125
Zhang S, Ma K, Chen L (2003) Response of photosynthetic
plasticity of Paeonia suffruticosa to changed light
environments. Environ Exper Bot 49:121-133
Zhao DQ, Hao ZJ, Tao J (2012) Effects of shade on plant
growth and flower quality in the herbaceous peony
(Paeonia lactiflora Pall.). Plant Physio Biochem 61:187-196
DOI:10.1016/j.plaphy.2012.10.005
Zhu H, Li X, Zhai W, Liu Y, Gao Q, Liu J, Ren L, Chen H,
Zhu Y (2017) Effects of low light on photosynthetic
properties, antioxidant enzyme activity, and anthocyanin
accumulation in purple pak-choi (Brassica campestris ssp.
Chinensis Makino). PLoS ONE 12:e0179305 DOI:10.1371/
journal.pone.0179305
Zhukov VI, Romanovskaja RN (1980) The length of the day
at accelerated growing spring wheat. Sci Tech Bul
Siberian Res Inst Plant Grow Breed 5:74-77