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Light source and CO2 concentration affect growth and anthocyanin content of
lettuce under controlled environment

Article in Horticulture, Environment and Biotechnology · December 2013

DOI: 10.1007/s13580-012-0821-9

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Hort. Environ. Biotechnol. 53(6):460-466. 2012.
DOI 10.1007/s13580-012-0821-9
Research Report

Light Source and CO2 Concentration Affect Growth and Anthocyanin

Content of Lettuce under Controlled Environment
1 1 1,2,3 1,2,3*
Yoo Gyeong Park , Ji Eun Park , Seung Jae Hwang , and Byoung Ryong Jeong
1
Department of Horticulture, Division of Applied Life Science (BK21 Program), Graduate School of
Gyeongsang National University, Jinju 660-701, Korea
2
Institute of Agriculture & Life Science, Gyeongsang National University, Jinju 660-701, Korea
3
Research Institute of Life Science, Gyeongsang National University, Jinju 660-701, Korea

*Corresponding author: [email protected]

Received June 13, 2012 / Revised September 24, 2012 / Accepted September 24, 2012
Ⓒ Korean Society for Horticultural Science and Springer 2012

Abstract. The effect of light source and CO2 concentration on the growth and anthocyanin content of lettuce
(Lactuca sativa L. ‘Seonhong Jeokchukmyeon’) grown in growth chambers was examined. The plant was grown
-2 -1
under 140 µmol･m ･s PPF provided by either cool white fluorescent lamps (F, the control), white (W) light emitting
diodes (LEDs), or a 8:1:1 mixture of red, blue and white (RBW) LEDs. Carbon dioxide concentration of the atmosphere
was maintained at either 350, 700, or 1,000 μmol･mol-1. The RBW treatment promoted vegetative growth of the shoot
and root. Chlorophyll fluorescence (Fv/Fm) was not significantly affected by the light source and CO2 concentration.
Total anthocyanin content of the plant supplied with 1,000 μmol･mol-1 CO2 was the greatest in the F treatment.
Photosynthetic rate significantly increased with the increasing CO2 concentration. These results suggested that the
RBW which provided a wider spectrum of PAR and the highest CO2 concentration provided the most the suitable
environment condition for vegetative growth of lettuce among the tested light sources. To obtain plants with even
higher quality, especially having greater content of anthocyanin, however, more considerations on supplemental light
source including white LED are necessary in terms of optimum intensity, photoperiod, and optimum ratios of mixing
with other LEDs.
Additional key words: closed system, fluorescent lamp, light emitting diode (LED), light quality, photosynthetic rate

Introduction available in the spectral range covering entire visible and

near-UV regions with a potential to supersede conventional
The plant factory in which crops are produced with lamps in efficiency (Tamulaitis et al., 2005). Combinations
optimization of the growth environment has potential of red (600-700 nm) and blue (400-500 nm) LEDs have
advantages of high productivity, year-round production, proven to be effective lighting sources for producing spinach,
pesticide-free harvest, lower water demand, and effective radish, and lettuce biomass in controlled environments (Goins,
land use (Kwon, 1996; Ono and Watanabe, 2006; Yeo, 2002; Goins and Yorio, 2000; Goins et al.,1997, 2001; Kim
2006). In a controlled environment, light is important for et al., 2006; Yorio et al., 2001). The combination of red and
plants not only as the energy source for photosynthesis but blue light was an effective lighting source to produce plant
also as a signal for development and morphogenesis. A biomass, and the addition of green light to blue and red light
major challenge to growing plants in the controlled system was also effective (Kim et al., 2004a, 2004b). Although the
will be controlling and supplying of a sufficient quantity and combination of red and blue LEDs has great potential for
quality of light (Bugbee and Salisbury, 1988; Kim et al., use as a light source to drive photosynthesis, plants are
2004c; Langhans and Dressen, 1988; Sager and Wheeler, adapted to utilize a wider spectrum of light to control photo-
1992; Salisbury and Bugbee, 1988). morphogenesis. Emerson and Rabinowitch (1960) discovered
Light emitting diodes (LEDs), in comparison with con- an enhancement of photosynthesis when two different light
ventional high-pressure sodium (HPS) and fluorescent lamps, beams, exciting two different pigments, by giving higher
are environment friendly and long-lifetime light sources rates than the sum of photosynthesis in the two beams given
 Hort. Environ. Biotechnol. 53(6):460-466. 2012. 461

-1
separately. nutrient solution (pH 6.5 and EC 1.5 mS･cm , Sonneveld
It is thought that such light sources as white LEDs may be and Straver, 1994) was supplied through a deep flow
more efficient for photosynthesis than those LEDs with just technique system.
one or two narrow bandwidth wavelengths (e.g. red, blue,
green, or a combination of red and blue), since white LEDs Light Source and CO2 Concentration Treatments
have some advantages of having a broad spectrum such as Light treatments examined in the present experiment were:
the sun light. Nowadays, high-power phosphor-converted cool white fluorescent lamps (F, the control) (Philips Co.,
white LEDs are commercially available, which are more Ltd., the Netherlands), white (W) light emitting diodes (LEDs)
energy efficient than incandescent lamps and are rapidly (FC Poibe Co., Ltd., Korea), and a 8:1:1 mixture in number
catching up with the energy performances of linear fluor- of LED chips (5 watt per LED chip) of red, blue and white
escent lamps (Pinho et al., 2007). It can be concluded that (RBW) LEDs (FC Poibe Co., Ltd., Korea) placed horizontally
white LEDs can be used as a light source to supplement 20-50 cm above the bench top in the controlled environment.
light for vegetables because it has many advantages of The average PPF measured with a digital photometer
polychromatic spectrum, high luminous efficiency, high light (HD2102.1, Delta OHM, Italy) at a 20 cm distance above
intensity, small size, long life span, easy operability and so the bench top before light source treatment were adjusted to
-2 -1
on (Hu et al., 2007). Despite these advantages of white LEDs, 140 ± 20 µmol･m ･s for each treatment. Light spectral
there have been only a few studies on the plant culture using distribution was scanned using a spectroradiometer (RPS-900R,
this light source. For practical plant factories, more studies International Light Co., USA) at the same 20 cm distance
are needed concerning the environmental conditions and above the bench top at an interval of 1 nm. The spectral
other factors, such as CO2 which is widely known to affect distribution and characteristics measured at three locations
plant growth and productivity (Gaastra, 1959), to produce within plant growing area in each light treatment are shown
the crop. In this study, three light sources, each with dif- in Fig. 1 and Table 1. Carbon dioxide concentration of the
ferent peak wavelengths and light quality, and CO2 con- atmosphere was maintained at either 350, 700, or 1,000
-1
centration were used to investigate their effect on growth μmol･mol during light cycles by adding pure CO2 from a
and anthocyanin content of lettuce, and to investigate the liquified CO2 tank. The plant factory was equipped with a
possibility of using white LEDs for the plant culture in plant
factories.

Materials and Methods

Plant Materials and Growth Conditions
Seeds of lettuce (Lactuca sativa L.) ‘Seonhong Jeokchuk-
myeon’ (Asia Seed Co., Korea) were sown in 240-cell plug
trays (60 cm × 41 cm × 5 cm) containing rockwool pellets
as the germination medium. Seedlings were grown in a
growth chamber (C1200H3, FC Poibe Co., Ltd., Korea) at
-2 -1
21℃ and under 140 ± 20 µmol･m ･s PPF provided by
fluorescent lamps (Philips Co., Ltd., the Netherlands). After
26 days, when the seedlings developed 3-4 true leaves
(approximately 4.2 cm in length), seedlings were transplanted
at a distance of 20 cm under respective light source and CO2
Fig. 1. Spectral distribution of light sources used in a controlled
concentration treatments. The culture conditions maintained environment experiment. The light sources used were F,
-2 -1
were 21 ± 2℃, 60 ± 10% RH, 140 ± 20 µmol･m ･s PPF, fluorescent lamp; W, white LED; and RBW, combined LED of
and a 12/12 hours (light/dark) photoperiod. A complete red, blue and white at 8:1:1.

Table 1. Spectral characteristics of fluorescent lamp (F), white (W) LEDs, and a mixture of red, blue and white (RBW) LEDs.

Photon flux (μmol･m-2･s-1)

Light source
300-400 nm 400-500 nm 500-600 nm 600-700 nm 700-800 nm
F 0.41 42.11 61.76 34.67 1.05
W 1.57 35.05 71.27 28.86 3.25
RBW 2.70 24.44 12.15 98.71 1.96
462 Yoo Gyeong Park, Ji Eun Park, Seung Jae Hwang, and Byoung Ryong Jeong

fan type CO2 distribution system inside to disperse the using a potable photosynthesis system (LI-6400, LI-COR,
injected CO2 throughout the plant factory. USA) immediately prior to harvesting. Measurement conditions
were as follows: leaf temperature, 20℃; flow rate, 500
-1 -2 -1
Measurements μmol･s ; PPF, 1,000 µmol･m ･s ; and CO2 concentration,
-1
Changes of fresh weights of the shoot and the root were 1,000 μmol･mol .
measured daily during the 20 days of culture after trans- The content of total anthocyanin was quantified from the
planting. Twenty days after initiating the light and CO2 absorbance at the wavelength of 530 nm using an UV
treatments, plant size, plant height, leaf length, leaf width, spectrophotometer (Uvikon 922, Kotron Instrumentals,
leaf area, number of leaves, leaf chlorophyll content, length Italy) after a 24 hours extraction at 4℃ in the dark after
of the longest root, chlorophyll fluorescence (Fv/Fm), fresh depositing 2 g leaf into 10 mL of 1.5 N HCl-95% ethanol
and dry weights of shoot and root, photosynthetic rate, and (85:15, v:v). The absorbance was calculated based on μg･
-1 -1
total anthocyanin content were measured. Plant size was g ･FW (Fuleki and Francis, 1968).
calculated as (plant height + plant width)/2. Leaf width was
taken on the youngest mature leaf. Leaf area was measured Experimental Design and Statistical Analysis
by using a leaf area meter (LI-3100, LI-COR Inc., USA). A completely randomized block design with 3 replications
For chlorophyll measurement, vigorously growing third leaf and 18 seedlings per treatment was employed in the ex-
from the top was extracted with an 80% (v/v) acetone periment. The treatment locations in a controlled environment
solution for 24 hours. Chlorophyll content was determined were randomly mixed between replications to minimize
by measuring absorbance by a spectrophotometer (Uvikon position effects. Data collected were analyzed for statistical
922, Kotron Instrumentals, Italy) at 645 and 663 nm, significance by the SAS (Statistical Analysis System, V.
according to the procedure described by Arnon (1949). 9.1, Cary, NC, USA) program. The experimental results
Chlorophyll fluorescence was measured in diurnal time were subjected to an analysis of variance (ANOVA) and
courses using a portable saturation-pulse fluorometer PAM- Duncan’s multiple range test. Graphing was performed with
2000 (Walz, Effeltrich, Germany). Measurements were taken Sigma Plot 10.0 (Systat Software, Inc., San Jose, CA, USA).
on the vigorously growing third leaf from the top. Saturation
pulses were applied to achieve a complete saturation of the Results and Discussion
PSII reaction centers. Dry weight was measured after drying
tissues for 72 hours in a drying oven (Model FO-450M, Jeio Growth of lettuce plants was significantly affected by
Technology Co., Ltd., Korea) at 80℃. light source and CO2 concentration (Table 2). Plant size
Photosynthesis, transpiration rate and stomatal conductance significantly increased with the increasing CO2 concen-
were measured 3 times per treatment. Photosynthesis was tration under the RBW, but not under the W. The plant grown
measured on the vigorously growing third leaf from the top under the RBW was the greatest with the rapid increase in

Table 2. Effect of light source and CO2 concentration on growth of lettuce measured at 46 days after sowing.
Light CO 2 Plant Leaf Leaf Leaf Length of Chlorophyll Fresh weight (g) Dry weight (g)
z -1 y No. of Chlorophyll
source (μmol･mol ) size length width area -1 longest fluorescence
2 leaves (μg·mg FW) Shoot Root Shoot Root
(A) (B) (cm) (cm) (cm) (cm ) root (cm) (Fv/Fm)
x
350 15.6 c 11.7 bc 8.1 c 334.4 d 6.0 a 0.63 ab 27.4 cd 0.82 a 8.3 c 0.69 d 0.58 ef 0.05 fg
F 700 15.6 c 9.2 d 7.8 d 273.6 d 5.0 b-d 0.62 ab 22.5 e 0.83 a 7.0 c 0.61 d 0.47 f 0.03 g
1,000 17.5 b 10.7 c 9.6 bc 298.7 d 5.6 ab 0.59 b 29.8 d 0.83 a 9.0 c 0.79 d 0.42 f 0.03 g
350 15.6 c 11.4 bc 8.7 c 462.3 c 5.3 b-d 0.62 ab 32.7 c 0.79 a 12.3 b 2.04 c 0.74 de 0.07 ef
W 700 15.6 c 11.1 bc 8.9 c 468.5 c 4.9 b-d 0.63 ab 30.5 c 0.81 a 13.5 b 2.26 c 0.67 d-f 0.08 de
1,000 16.0 bc 11.0 bc 9.0 c 479.5 c 4.9 cd 0.65 a 39.4 ab 0.81 a 24.8 a 4.64 b 1.00 bc 0.13 ab
350 17.5 b 12.1 ab 9.6 bc 585.9 b 5.6 ab 0.59 b 36.7 b 0.80 a 23.5 a 5.37 ab 1.24 ab 0.14 a
RBW 700 19.5 a 13.0 a 10.1 b 648.1 ab 5.5 a-c 0.54 c 39.4 ab 0.81 a 24.8 a 4.64 b 1.00 bc 0.13 ab
1,000 21.0 a 12.1 ab 10.9 a 664.5 a 5.1 b-d 0.63 ab 42.4 a 0.82 a 26.4 a 5.65 a 1.35 a 0.15 a
A *** NS *** *** NS ** *** NS *** *** *** ***
F-test B NS *** NS NS *** * ** NS NS NS * NS
A*B ** ** ** NS NS NS ** NS * NS NS NS
z
The light sources used were F, fluorescent lamp; W, white LED; and RBW, combined LED of red, blue and white at 8:1:1.
y
(Plant height + plant width)/2.
x
Mean separation within columns by Duncan’s multiple range test at P = 0.05.
NS,*,**,***
Nonsignificant or significantly different at P = 0.05, 0.01, and 0.001, respectively.
 Hort. Environ. Biotechnol. 53(6):460-466. 2012. 463

Fig. 3. Effect of light source and CO2 concentration on growth of

lettuce measured at 46 days after sowing. The light sources used
were F, fluorescent lamp; W, white LED; and RBW, combined
LED of red, blue and white at 8:1:1.

-1
1,000 μmol･mol CO2 as compared to the control. Chlorophyll
concentration was the greatest when plants were grown
-1
under the W and were applied with 1,000 μmol･mol CO2,
and the least when plants were grown under the RBW and
-1
were applied with 700 μmol･mol CO2. The length of the
longest root of the plants grown under the RBW increased
by 54% as compared to the control with increasing CO2
-1
concentration with the applied 1,000 μmol･mol CO2 (Table
2). Chlorophyll fluorescence (Fv/Fm) was not significantly
affected by light source and CO2 concentration. Fresh weights
of shoot and root in the control was not significantly affected
Fig. 2. Effect of light source and CO2 concentration on changes by CO2 concentration. Fresh and dry weight of root of the
of fresh weight of lettuce grown in a controlled environment
-1 -1
plants grown under the RBW was the greatest with the
supplemented with CO2 at A, 350 μmol･mol ; B, 700 μmol･mol ; applied CO2 at 1,000 μmol･mol .
-1
-1
or C, 1,000 μmol･mol . The light sources used were F, fluor-
escent lamp; W, white LED; and RBW, combined LED of red, Thus, the RBW treatment promoted overall vegetative
blue and white at 8:1:1. Data represent means ± SE of 3 growth, such as plant size and leaf growth, as compared to
measurements. the control (Fig. 3). Plant leaves readily absorb red and blue
light, so absorbance is high and reflection is relatively low
fresh weight (Fig. 2). The leaf length was the greatest when in these ranges of the spectrum (Kim et al., 2004a; Klein,
the plant was grown under the RBW. The leaf width 1992; Smith, 1993). The peak emissions of the blue and red
increased with increasing CO2 concentration under the RBW, LEDs coincide closely with the absorption peaks of chloro-
-1
and were applied with 1,000 μmol･mol CO2. The leaf phyll a and b, and the reported wavelengths are of maximum
width increased by 34% when plants were grown under the photosynthetic efficiency (McCree, 1972). Light is absorbed
-1
RBW and applied with 1,000 μmol･mol CO2 as compared by phytochrome and cryptochrome photo-receptors, which
-1
to the F and applied with 350 μmol･mol CO2. The leaf promote the expression of genes that, change the fate of the
width was the greatest when plants were grown under the shoot apical meristem from vegetative growth to reproductive
RBW. The leaf area increased by 60 and 122%, respectively, development (Amasino, 1996). Red spectrum (600-700 nm)
when plants were grown under the W and RBW and were of the RBW (a mixture of red, blue and white LEDs at 8:1:1)
-1
applied with 1,000 μmol･mol CO2 as compared to the F used in this study has the greatest photon flux, which is
-1
and applied with 1,000 μmol･mol CO2. The number of about 3 times greater as compared to the F and W treatments
leaves decreased by 19% and 15%, respectively, when plants (Table 1). Mor et al. (1980) suggested that red light was
were grown under the W and RBW, and were applied with most effective in increasing the transport of assimilates to
464 Yoo Gyeong Park, Ji Eun Park, Seung Jae Hwang, and Byoung Ryong Jeong

Fig. 5. Effect of light source and CO2 concentration on total antho-

cyanin content of lettuce measured at 46 days after sowing. The
light sources used were F, fluorescent lamp; W, white LED; and
RBW, combined LED of red, blue and white at 8:1:1. Data
represent means of 3 measurements.

plants grown under the W (13.11 μmol CO2･m-2･s-1) and

showed similar results with plants grown under the F and
RBW. At 1,000 μmol･mol-1 CO2, no significant difference
was shown among light source treatments, and the photo-
synthetic rate was 14.17 μmol CO2･m-2 ･s-1 (Fig. 4). Trans-
piration rate (0.11 to 0.37 μmol H2O･m-2･s-1) significantly
increased with increasing CO2 concentration under the F.
Transpiration rate of the plants grown under the W was the
least when fed with 350 μmol･mol-1 CO2. Transpiration rate
and stomatal conductance (Fig. 4) were the greatest in plants
grown under the RBW and applied with 1,000 μmol･mol-1
CO2. Similar results were reported by Yorio et al. (2001)
who described that stomatal opening was stimulated, but
photosynthesis was not enhanced, in leaves of lettuce under
red-LED light supplemented with blue light.
Fig. 4. Effect of light source and CO2 concentration on photo- Total anthocyanin content in the leaf grown under the
synthesis, transpiration rate and stomatal conductance of lettuce control tended to increase with increasing CO2 concen-
measured at 46 days after sowing. The light sources used were
F, fluorescent lamp; W, white LED; and RBW, combined LED tration, but not under the W and RBW (Fig. 5). Total
of red, blue and white at 8:1:1. Data represent means of 3 anthocyanin content per unit fresh weight was not sig-
measurements. nificantly affected by the CO2 treatment. Total anthocyanin
content was the greatest with the plants grown under the F
the shoot tips of rose plants, and it promoted shoot sink and when applied with 1,000 μmol･mol-1 CO2. The W (3.25
activity by increasing the unloading process. μmol･m-2･s-1) and RBW (1.96 μmol･m-2･s-1) had absolute
Photosynthetic rate significantly increased with increasing irradiance in the far-red (700-800 nm) range, and as a result
CO2 concentration in all light source treatments (Fig. 4). anthocyanin content in those treatments was lower than in
However, photosynthetic rate measured in this experiment the F (1.05 μmol･m-2･s-1) (Table 1). There is also evidence
was not affected by the light source. The reason for this is reported for the involvement of phytochromes in anthocyanins
not clear, but may be due to altered photosynthetic rates as synthesis (Alokam et al., 2002; Ramalho et al., 2002; Yanovsky
affected by time of measurement, since all samples were et al., 1998). When Li and Kubota (2009) reported that the
measured on the same date but at quite different times. At supplemental far-red treatment (red/far-red ratio = 0.5)
-1
350 μmol･mol CO2, photosynthetic rate of plants grown significantly reduced anthocyanin concentration as compared
under the F was higher than in the other treatments. At 700 to the control (red/far-red ratio = 11.5), while the supplemental
-1
μmol･mol CO2, photosynthetic rate was the greatest in red treatment had no significant effects, despite its high
 Hort. Environ. Biotechnol. 53(6):460-466. 2012. 465

red/far-red ratio of 38.5. The phytochrome response may be carbon dioxide, temperature and stomatal diffusion resistance.
already saturated under the control and the supplemental red Mededlinger van de Landbouwhogeschool Te Wagenigen 59:1-68.
Goins, G.D. 2002. Growth, stomatal conductance, and leaf surface
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