www.ccsenet.org/jas Journal of Agricultural Science Vol. 4, No.

4; 2012

Effects of Different Light Sources on the Growth of Non-heading

Chinese Cabbage (Brassica campestris L.)

Huimin Li
College of Agronomy, Nanjing Agricultural University. Nanjing, Jiangsu, 210095, China
E-mail: [email protected]

Canming Tang (Corresponding author)

College of Agronomy, Nanjing Agricultural University. Nanjing, Jiangsu, 210095, China
Tel: 86-258-486-0062 E-mail: [email protected]

Zhigang Xu (Corresponding author)

College of Agronomy, Nanjing Agricultural University. Nanjing, Jiangsu, 210095, China
E-mail: [email protected]

Xiaoying Liu
College of Agronomy, Nanjing Agricultural University. Nanjing, Jiangsu, 210095, China
E-mail: [email protected]

Xuelin Han
Kingsun, optoelectronic co.,ltd. Dongguan, Guangdong, 523565, China
E-mail: [email protected]

Received: December 23, 2011 Accepted: January 4, 2011 Online Published: February 2, 2012
doi:10.5539/jas.v4n4p262 URL: http://dx.doi.org/10.5539/jas.v4n4p262

This research is supported by National Natural Science Fundation (30972035), the National 863 Plans Program
(2011AA03A1), the National and Jiangsu Science and Technology Key Program (2011BAE01B00) and
(BE2011197) .

Abstract
To date, little is known about the effects of different light sources on the growth and quality of non-heading
Chinese cabbage (Brassica campestris L.). The objective of present study was to evaluate the effects of
light-emitting diodes (LEDs) light sources (blue, blue plus red, red), fluorescent lamps and sunlight on growth
and vitamin C, soluble protein, sucrose, soluble sugar, starch and pigment concentrations in non-heading Chinese
cabbage seedlings. The dry mass of shoots and the fresh and dry masses of roots were highest in seedlings grown
under red LEDs with weak lights. The fresh mass of roots and starch concentration were highest under red LEDs
despite of the altered photosynthetic photo flux density (PPFD) levels. The concentrations of chlorophylls and
vitamin C were greatest under blue LEDs with altered PPFD levels. The numbers of flower buds and open
flowers were highest under red LEDs and blue plus red LEDs and were higher under LEDs than fluorescent
lamps. The duration of flowering was highest under red LEDs and blue plus red LEDs. The present results
demonstrate that LED light sources are more effective than fluorescent lamps for vegetative and reproductive
growth of non-heading Chinese cabbage. Moreover, blue LEDs benefit vegetative growth, while red LEDs and

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blue plus red LEDs support reproductive growth in non-heading Chinese cabbage. In the artificial cultivation and
subsequent transplanting of the life cycle of plants, the light source can be selected to meet the requirements of
different growth stages of plants and be used to promote the subsequent process in the industrial production of
non-heading Chinese cabbage.
Keywords: Light-emitting diodes (LEDs), Non-heading Chinese cabbage, Vitamin C, Starch, Pigment, Flower
bud
1. Introduction
Light plays a key role in plant life, determining their photo-morphogenesis and photosynthesis rate (Avercheva et
al., 2009). The sun emits the most of its radiation in the visible range, it covers the range of wavelength from
400-700nm (Kolawole, et al., 2010). The integration, quality, duration and intensity of red, far-red, blue, UV-A
(320–500 nm) and UV-B (280–320 nm) light have a profound influence on plants by triggering physiological
reactions to control their growth and development (Briggs et al., 2001; Briggs and Olney, 2001; Clouse, 2001).
LEDs are solid-state, long-lasting and durable sources of narrow-band light that can be used in a variety of
horticultural and photo-biological applications (Stutte, 2009), including controlled research environments
(Avercheva et al., 2009), lighting for tissue culture (Li et al., 2010) and supplemental and photoperiod lighting
for greenhouses (Morrow, 2008). Because of their potential to be implemented in dynamic lighting strategies to
control plant growth, development, physiological responses and production, it is important to learn more about
the influence of light quality on these processes (Folta and Childers, 2008; Lefsrud et al., 2008; Massa et al.,
2008).
Various studies have shown that LEDs have been successfully used for cultivation in several horticultural plant
species, including lettuce (Bula et al., 1991; Hoenecke et al., 1992; Yanagi et al., 1996; Okamoto et al., 1997;
Yorio et al., 2001; Kim et al., 2004; Kim et al., 2006; Stutte et al., 2009; Li and Kubota, 2009), cucumber
(Menard et al., 2006; Brazaityte et al., 2009), pepper (Brown et al., 1995; Schuerger et al., 1997), spinach (Yorio
et al., 2001), radish (Yorio et al., 2001; Tamulaitis et al., 2005), Chinese cabbage (Avercheva et al., 2009) and
tomato (Kaneko-Ohashi et al., 2004; Menard et al., 2006; Brazaityte et al., 2010; Liu et al., 2011). Although
previous studies have identified various physiological and morphological effects of light quality in many plant
species, few reports have addressed the effect of LED light sources, sunlight and fluorescent lamps on the
growth of non-heading Chinese cabbage (Brassica campestris L.). Non-heading Chinese cabbage originated
from China and has a long cultivation history. Its leaves contain many beneficial materials, and it is an important
cultivated vegetable species in China (Hu and Hou, 2010). Variations in light conditions will affect the metabolic
processes such as growth and yield (Jaimez and Rada, 2011). The objective of the present study was to examine
the effects of blue LEDs, red LEDs, blue plus red LEDs (B:R=1:8), fluorescent lamps and sunlight on the growth
and quality of non-heading Chinese cabbage seedlings at different stages of development and to select the best
light sources for the cultivation of seedlings under a controlled environment.
2. Materials and Methods
2.1 Plant Materials
The experiments were performed in a greenhouse at Nanjing Agricultural University with non-heading Chinese
cabbage (Brassica campestris L.) cultivar 605. Seeds with a similar size were selected for sowing. Seeds were
sown in cells filled with vermiculite and peat (1:1 by volume) for cultivation, with one seed per cell. After seven
days, seedlings with two expanded cotyledons were transferred to the different lights.
2.2 Light Treatments
Seedlings were grown under red light-emitting diodes (LEDs), blue LEDs, a mixture of blue plus red LEDs
(B:R=1:8) and fluorescent lamps. The growth temperature was set at 24–26°C , and the relative humidity
fluctuated between 40 and 50%. The photoperiod was 12 hours. Seedlings were randomly assigned to each light
treatment, and LEDs arrays were randomly assigned positions in the greenhouse. Seedlings of the first group
were kept at a photosynthetic photo flux density (PPFD) of 80 μ mol m–2s–1 (weak light) for 60 days (60 days).
Seedlings of the second group were first grown at weak light for 30 days, then transferred to sunlight (normal
level of 350 μ mol m–2s–1) at the 31st day and cultured for 30 days (60 days). Seedlings of the third group were
first grown at weak light for 30 days, then transferred to a normal level of sunlight on the 31st day and cultured
for 30 days, and then transferred to weak light on the 61st day until flowering (90, 120 days). Numbering the
flower buds and open flowers was from 90th to 120th days. Each group had 30 seedlings. Each experiment was
replicated three times. The spectral energy distribution of the blue plus red LEDs, blue LEDs, red LEDs, sunlight
and fluorescent lamps was measured using an OPT-2000 spectral photometer (Optpeco, Beijing, China). FL:

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Fluorescent lamps. B+R: 11.1% blue plus 88.9% red LEDs. B: Blue LEDs. R: Red LEDs. a: Growth under LEDs
and FL at a PPFD level of 80 μmol m–2s–1 (weak light) for 60 days. b: Iinitial growth at a PPFD level of 80 μmol
m–2s–1 for 30 days followed by sunlight for 30 days. c: Initial growth at a PPFD level of 80 μmol m–2s–1 for 30
days followed by sunlight for 30 days and then weak light at 61 days until flowering.
(1) The blue plus red LEDs array (B+R) was supplied with 11.1% blue light with peak emission at 460 nm and
88.9% red light with peak emission at 660 nm (20 nm band-width at half-peak height).
(2) The blue LEDs array (B) was supplied with 100% blue light with peak emission at 460 nm (20 nm
band-width at half-peak height).
(3) The red LEDs array (R) was supplied with 100% red light with peak emission at 660 nm (20 nm band-width
at half-peak height).
(4) The fluorescent lamp array (FL) was supplied a broad spectrum of light with peak emission at 400–700 nm.
(5) The sunlight light array (SUN) was supplied a mean PPFD of 350 μ mol m–2s–1 with peak emission at
380–2600 nm.
2.3 Growth Measurements
Seedlings were destructively sampled after 60 (the first and second group) and 90 (the third group) days of
growth. To determine dry mass, the seedlings were dried at 85°C until a constant mass was reached. The mass of
each seedling was measured using an electronic balance.
For 2.4, 2.5, 2.6 and 2.7 measurements described below, seedlings were all destructively sampled after 60 (the
first and second group) and 90 (the third group) days of growth. After sampling, leaves were weighed as
required.
2.4 Pigment Measurements
Leaves were weighed to 0.1 g (fresh weight, W), and 10 ml (V) of 80% acetone was added to 0.1 g of leaf
samples placed into a mortar with quartz sand. The chlorophyll was extracted until the leaf turned white. The
optical density (OD) was measured with a UV-1200 spectrophotometer (Jin Peng, Shanghai, China) at 470 nm
for carotenoid (OD470), at 663 nm for chlorophyll a (OD663), and at 645 nm for chlorophyll b (OD645). The
concentrations of chlorophyll a, chlorophyll b and chlorophyll (a+b) were determined using the following
equations (Lichtenthaler and Wellburn, 1983):
Chlorophyll a (mg / g) = (12.72 OD663－2.59 OD645) V / 1000 W
Chlorophyll b (mg / g) = (22.88 OD645－4.67 OD663) V / 1000 W
Chlorophyll (a＋b) (mg / g) = (8.05 OD663 + 20.29 OD645) V / 1000 W
Carotenoid (mg / g) = (1000 OD470－3.27 Ca－104 Cb) V / (229×1000 W)
Where V is the total volume of acetone extract (ml), W is the fresh weight (g) of the sample, Ca is the
concentration of chlorophyll a, and Cb is the concentration of chlorophyll b (Li et al., 2010).
2.5 Soluble Protein Measurements
Leaves (1.0 g of fresh weight, W) were ground in a mortar with liquid nitrogen, to which 5 ml (V1) of 0.067
mol/l potassium phosphate buffer (PBS) was added, and were then filtered through filter paper. The extract was
centrifuged at 12,000 g for 10 min, and the supernatant was removed. The extract (1 ml, V2) and Coomassie
brilliant blue G-250 (5 ml) was thoroughly mixed. The optical density was measured using a UV-1200
spectrophotometer at 595 nm. To determine a standard curve, 0, 0.2, 0.4, 0.6, 0.8, and 1 ml of 100 μg/l bovine
serum albumin was added to 6 volumetric flasks, and distilled water was added to reach a volume of 1 ml. The
optical density was measured by a UV-1200 spectrophotometer at 595 nm (ρ). The concentration of soluble
protein was determined using the following equation: soluble protein (mg / g) = ρ V1 / W V2 (Li et al., 2010).
2.6 Vitamin C Measurements
Leaves of (1.0 g, fresh weight, W) were ground in a mortar with liquid nitrogen. Next, 5 ml (V1) of 5%
trichloroacetic acid (TCA) was added and the mixture was filtered through filter paper. The extract was
centrifuged at 10,000 g for 10 min, and the supernatant was removed. The extract (1.0 ml, V2) and 1.0 ml of
ethanol were thoroughly mixed. Next, 0.5 ml of 0.4% phosphoric acid-ethanol, 1 ml of 0.5% 1, 10-
phenanthroline-ethanol and 0.5 ml of 0.03 g/l ferric chloride were added for a total volume of 5 ml. The optical
density was measured using a UV-1200 spectrophotometer (Jin Peng, Shanghai, China) at 534 nm. To obtain a
standard curve, 0, 0.2, 0.4, 0.6, 0.8, or 1 ml of 100 mg/l bovine serum albumin was added to 6 volumetric flasks,

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and distilled water was added to reach a volume of 1 ml. The optical density was measured by a UV-1200
spectrophotometer at 534 nm (ρ). The concentration of vitamin C was determined using the following equation:
vitamin C concentration (mg / g) = ρ V1 / W V2 (Li et al., 2010).
2.7 Sugar and Starch Measurements
Leaves (0.5 g, dry weight) were ground in a mortar with liquid nitrogen. Then 1 ml of 80% ethanol was added,
and the mixture was filtered through filter paper. The filtrates were recovered, and the residues were washed
again with 70% ethanol and filtered. Both filtrates were mixed, and 3 ml of distilled water was added. The
extract was centrifuged at 12,000 g for 15 min, and 1 ml of supernatant was collected. Soluble sugar
concentration was determined by the sulfuric acid-anthrone method and measured at 620 nm. Sucrose
concentration was determined using the phloroglucinol method and measured at 480 nm (Zhang, 2009).
Takahashi’s method was used for starch extraction (Takahashi et al., 1995). The residue obtained after ethanol
extraction was resuspended with 0.1 mol/l sodium acetate buffer (pH 4.8) and boiled for 20 min. The gelatinized
starch was digested with amyloglucosidase for four hours at 37°C and boiled again to stop the enzymatic
reaction. After cooling, the mixture was centrifuged, and the amount of soluble sugar in the supernatant was
determined by anthrone colorimetry (Li et al., 2010). The starch concentration was estimated by converting
glucose to starch equivalents using a factor of 0.9.
2.8 Statistical Analysis
Statistical analyses were conducted with Statistical Product and Service Solutions (SPSS) for Windows,
Version 16.0 (SPSS, Japan). Data were analyzed using analysis of variance (ANOVA), and the differences
between means were tested using Tukey’s Test (P＜0.05).
3. Results
3.1 The Growth of Non-heading Chinese Cabbage
The effects of different light sources on growth of non-heading Chinese cabbage seedlings under a PPFD of 80
μmol m–2s–1 (weak light) for 60 days varied significantly (Table 1). The fresh mass of shoots was higher in
seedlings grown under blue + red, blue and red LEDs and lower under fluorescent lamps. The dry mass of shoots
and those of fresh and dry roots were greatest in seedlings under red LEDs and lowest under fluorescent lamps,
an indication that LEDs light sources are more suitable for the growth of non-heading Chinese cabbage than
fluorescent lamps and that red LEDs benefit biomass accumulation in non-heading Chinese cabbage.
Different light sources had variable effects on growth of non-heading Chinese cabbage seedlings that were first
grown under weak light for 30 days and then transferred to sunlight for 30 days (Table 2). The dry masses of
shoots and roots were greatest in seedlings under B:R=1:8 LEDs and lowest under fluorescent lamps. The fresh
mass of shoots and roots were greatest in seedlings under red LEDs and lowest under fluorescent lamps. This is
an indication that B:R=1:8 LEDs benefit the accumulation of dry biomass. The after-effects of different light
sources on non-heading Chinese cabbage seedlings were obvious.
Different light sources had variable effects on growth of non-heading Chinese cabbage seedlings before
flowering (Table 3). The fresh mass of shoots was greatest in seedlings under blue LEDs, whereas the fresh and
dry masses of roots were greatest in seedlings under red LEDs and lowest under fluorescent lamps. For the shoot
dry mass, red LEDs had similar higher; while fluorescent lamps gave the least shoot dry mass than any of the
lights sources examined.
3.2 The Concentrations of Pigments
The leaf pigments of non-heading Chinese cabbage seedlings varied in response to the weak light. The leaf
pigments also varied when they were first grown at weak light and then transferred to sunlight. The
concentrations of chlorophyll a, chlorophyll b, total chlorophyll and carotenoid were greatest in seedlings under
blue LEDs (Figures 1a & b).
Different light sources had variable effects on the concentrations of pigments in non-heading Chinese cabbage
seedlings before blooming. The concentrations of chlorophyll a, chlorophyll b and total chlorophyll were greatest
in seedlings under blue LEDs and lowest in seedlings under fluorescent lamps (Figure c).
3.3 Vitamin C Concentration
The concentration of vitamin C was greatest in non-heading Chinese cabbage seedlings under blue plus red
LEDs with weak light, followed by those under blue LEDs and lowest in seedlings under fluorescent lamps
(Figure 2). The seedlings adapted to new illumination conditions with altered PPF levels. The concentration of

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vitamin C was greatest in seedlings under blue LEDs and lowest in seedlings under fluorescent lamps with
altered PPF levels.
3.4 Soluble Protein Concentration
The concentration of soluble protein was highest under red LEDs in seedlings with the weak light, followed by
fluorescent lamps and blue LEDs and lowest under blue plus red LEDs (Figure 3). The seedlings adapted to new
illumination conditions with altered PPF levels. The concentration of soluble protein was greatest in seedlings
under blue LEDs and blue plus red LEDs with altered PPF levels.
3.5 Sugar and Starch Concentrations
The sugar and starch concentrations of seedlings varied in response to different weak lights. Both concentrations
also varied when seedlings were first grown at weak light and then transferred to sunlight. The concentrations of
sucrose and soluble sugar were greatest in seedlings under blue LEDs, followed by those under red LEDs and
blue plus red LEDs and lowest in seedlings under fluorescent lamps (Figures 4a & b). The starch concentration
was greatest in seedlings under red LEDs.
The sugar and starch concentrations of seedlings varied in response to different lights before blooming. The
concentrations of sucrose, soluble sugar and starch were greatest in seedlings grown under red LEDs, followed
by those under blue plus red LEDs and lowest in seedlings under fluorescent lamps (Figure 4c). These results
revealed that red LEDs are the best light source for accumulation of sucrose, starch and soluble sugar in
non-heading Chinese cabbage.
3.6 The Number of Flower Buds and the Duration of Flowering
Different light sources had variable effects on the development of flowers in non-heading Chinese cabbage
seedlings from 90 to 120 days (Table 4). The number of open flowers was highest in seedlings under red LEDs
and blue plus red LEDs and the number of flower buds was higher in seedlings under LEDs than in those under
fluorescent lamps. The duration of flowering was longest in seedlings under red LEDs and blue plus red LEDs.
4. Discussion and Conclusion
4.1 Red LEDs Are the Best Light for Accumulation of Dry Mass and Photosynthates in Plants
The present study demonstrated that red LEDs benefits the dry biomass accumulation of non-heading Chinese
cabbage (Tables 1, 2 &3). However, the present study are inconsistent with those previous studies (Brown et al.,
1995; Goins et al., 1997; Okamoto et al., 1997; Yorio et al., 2001; Kim et al., 2004; Avercheva et al., 2009) that
showed the best light sources for dry biomass accumulation are related to species or cultivars. LEDs lights show
a significant superiority on plant growth and biomass. Red LEDs has variable effects on different plant species.
Light quality regulates the carbohydrate metabolism of higher plants, and carbohydrate content is increased
under red light. The accumulation of starch in chloroplasts, which is enhanced by red light, may cause inhibition
of photosynthesis (Kowallik et al., 1982). Red light may inhibit the translocation process of photosynthates
(Saebo et al., 1995). The excess starch accumulation counters leaf photosynthesis (Bondada and Syvertsen,
2005). Red light enhances starch accumulation in Glycine and Sorghum species, the application of blue light
sources is a means of studying the regulation of photosynthetic carbon metabolism in relation to plant growth
(Britz and Sager, 1990). The present study, which is consistent with previous studies, reveal that the starch
concentration is greatest in seedlings grown under red LEDs with altered PPFD levels (Figure 4) and red LEDs
are advantageous to accumulation of starch in non-heading Chinese cabbage, moreover, the red LEDs benefits
the accumulation of dry biomass in non-heading Chinese cabbage (Tables 1, 2 & 3). Red LEDs may promote
accumulation of the photosynthetic products and the dry biomass but inhibit the translocation of photosynthetic
products out of leaves; thus, the starch ultimately accumulated in leaves. Red LEDs may be used as the main
light sources for reproductive growth of non-heading Chinese cabbage. In the artificial cultivation of plants, red
LEDs may be the preferred light source, to get achieve a larger amount of dry matter and yield.
4.2 The Effects of Blue LEDs on the Accumulation of Chlorophylls and the Quality of Plants
The present results indicated that blue LEDs light is beneficial to pigment accumulation. The present results are
consistent with previous studies (Senger, 1982; Saebo et al., 1995; Tanaka et al., 1998; Poudel et al., 2008;
Kurilcik et al., 2008). Moreover, the blue LEDs play a key role in accumulation of chlorophylls.
The present results indicated that blue LEDs benefits vitamin C, sucrose and soluble sugar accumulation and
nutritional quality of non-heading Chinese cabbage seedlings. LEDs light is beneficial to soluble protein
accumulation and to nutritional quality of seedlings. The results in the present study demonstrate that blue LEDs
benefits the accumulation of vitamin C and soluble protein, which are consistent with reports by Yang et al.

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(2010) and Zhang et al. (2010). However, the soluble sugar concentration was greatest under blue LEDs (Figure
4), which is inconsistent with the results of Yang et al. (2010) and Zhang et al. (2010). Blue LEDs benefits the
accumulation of nutritional substances, and these effects may correlate with plant species or cultivars. The
nutritional quality of plants could be varied by selecting special light sources under controlled growth
environments. For the purpose of improving nutritional quality of vegetables, blue LEDs should be chosen as the
preferred light source in artificial cultivation and subsequent transplanting of plants.
4.3 Red LEDs Are the Best Light for Flowering
Spectral quality has a major influence on induction rate of flower bud and subsequent development. The
numbers of Cyclamen flower buds and open flowers were highest in plants grown under a mixture of blue plus
red LEDs (B:R=10) compared with fluorescent lamps and other light sources (Heo et al., 2003). However, the
development of visible flower buds in marigolds was about five times greater in fluorescent lamps than in blue
or red LEDs (Heo et al., 2002). Monochromatic blue light delayed flowering in Arabidopsis possibly by
influencing cryptochromes (Mockler et al., 1999). The present study showed that the number of flowers was
highest in seedlings grown under red LEDs and blue plus red LEDs, and the number of flower buds was higher
in seedlings grown under LEDs than fluorescent lamps (Table 4). Red LEDs and blue plus red LEDs are more
suitable for flower opening in non-heading Chinese cabbage. The findings from the present study are consistent
with those of Heo et al. (2003) and Mockler et al. (1999), but inconsistent with a report from Heo (2002). The
shift in plants from vegetative growth to floral development is regulated by red-far-red light receptors
(phytochromes) and blue-ultraviolet A light receptors (cryptochromes) (Guo et al., 1998). The present study also
showed that the concentrations of sucrose and starch were greatest in seedlings grown under red LEDs before
blooming and red LEDs benefitted dry biomass accumulation in non-heading Chinese cabbage. The number of
flower buds and open flowers and the duration of flowering may correlate with the dry biomass and
photosynthates, it regulated by different light receptors.
In conclusion, dry biomass and starch of non-heading Chinese cabbage accumulate under red LEDs light sources.
In addition, red LEDs also benefits the flower opening. The pigment and nutritional substances (vitamin C,
soluble sugar and soluble protein) accumulate under blue LEDs light sources. Red LEDs should be selected as
the preferred light source in the artificial cultivation and subsequent transplanting of plants to obtain a higher
biomass and more flowers. By contrast, blue LEDs should be used as the preferred light source for higher
nutritional quality to improve the growth and development of plants.
References
Avercheva, O.V., Berkovich, Y.A., Erokhin, A.N., Zhigalova, T.V., Pogosyan, S.I., & Smolyanina, S.O. (2009).
Growth and photosynthesis of Chinese cabbage plants grown under light-emitting diode-based light source.
Russian Journal of Plant Physiology, 56, 14-21. http://dx.doi.org/10.1134/S1021443709010038
Bondada, B.R., & Syvertsen, J.P. (2005). Concurrent changes in net CO2 assimilation and chloroplast
ultrastructure in nitrogen deficient citrus leaves. Environmental and Experimental Botany, 54, 41-48.
http://dx.doi.org/10.1016/j.envexpbot.2004.05.005.
Brazaityte, A., Duchovskis, P., Urbonaviciute, A., Samuoliene, G., Jankauskiene, J., Kasiuleviciute-Bonakere, A.,
Bliznikas, Z., Novickovas, A., Breive, K., & Zukauskas, A. (2009). The effect of light-emitting diodes lighting
on the growth of cucumber transplants and after-effect on yield. Zemdirbyste Agriculture, 96, 102-118.
http://zemdirbyste-agriculture.lzi.lt/96(3)tomas/96_3_tomas_102_118.pdf
Brazaityte, A., Duchovskis, P., Urbonaviciute, A., Samuoliene, G., Jankauskiene, J., Sakalauskaite, J.,
Sabajeviene, G., Sirtautas, R., & Novickovas, A. (2010). The effect of light-emitting diodes lighting on the
growth of tomato transplants. Zemdirbyste Agriculture, 97, 89-98.
http://www.lzi.lt/tomai/97(2)tomas/97_2_tomas_str10.pdf
Briggs, W.R., Beck, C.F., Cashmore, A.R., Christie, J.M., & Hunghes, J. (2001). The phototropin family of
photoreceptors. Plant Cell, 13, 993-997. http://dx.doi.org/10.1105/tpc.13.5.993
Briggs, W.R., & Olney, M.A. (2001). Photoreceptors in plant photomorphogenesis to date, five photochromes,
two cryptochrome, one phototropin and one superchrome. Plant Physiology, 25, 85-88.
http://dx.doi.org/10.1104/pp.125.1.85
Britz, S.J., & Sage, J.C. (1990). Photomorphogenesis and photoassimilation in soybean and sorghum grown under
broad spectrum or blue-deficient light sources. Plant Physiology, 94, 448-454.
http://www.jstor.org/stable/4273112

Published by Canadian Center of Science and Education 267

www.ccsenet.org/jas Journal of Agricultural Science Vol. 4, No. 4; 2012

Brown, C.S., Schuerger, A.C., & Sager, J.C. (1995). Growth and photomorphogenesis of pepper plants under red
light-emitting diodes with supplemental blue or far-red lighting. Journal of the American Society of Horticultural
Science, 120, 808-813. http://www.ncbi.nlm.nih.gov/pubmed/11540133
Bula, R.J., Morrow, R.C., Tibbitts, T.W., Barta, D.J., Ignatius, R.W., & Martin, T.S. (1991). Light-emitting diodes
as a radiation source for plants. HortScience, 26, 203-205. http://www.ncbi.nlm.nih.gov/pubmed/11537727
Clouse, S.D. (2001). Integration of light and brassinosteroid signals in etiolated seedling growth. Trends in Plant
Science, 6, 443-445. http://dx.doi.org/10.1016/S1360-1385(01)02102-1
Folta, K.M., & Childers, K.S. (2008). Light as a growth regulator: controlling plant biology with
narrow-bandwidth solid-state lighting systems. HortScience, 43, 1957-1964.
http://hortsci.ashspublications.org/content/43/7/1957.full
Goins, G.D., Yorio, N.C., Sanwo, M.M., Brown, C.S., & Sager, J.C. (1997). Photomorphogenesis, photosynthesis,
and seed yield of wheat plants grown under red light-emitting diodes (LEDs) with and without supplemental
blue lighting. Journal of experimental botany, 48, 1407-1413. http://dx.doi.org/10.1093/jxb/48.7.1407
Guo, H., Yang, H., Mockler, T.C., & Lin, C. (1998). Regulation of flowering time by Arabidopsis photoreceptors.
Science, 279, 1360-1363. http://dx.doi.org/10.1126/science.279.5355.1360
Heo, J., Lee, C., Chakrabarty, D., & Paek, K. (2002). Growth responses of marigold and salvia bedding plants as
affected by monochromic or mixture radiation provided by a light emitting diode (LED). Plant Growth
Regulation, 38, 225-230. http://dx.doi.org/10.1023/A:1021523832488
Heo, J., Lee, C., Chakrabarty, D., & Paek, K. (2003). Influence of light quality and photoperiod on flowering of
Cyclamen persicum Mill. cv. ‘Dixie White’. Plant Growth Regulation, 40, 7-10.
http://dx.doi.org/10.1023/A:1023096909497
Hoenecke, M.E., Bula, R.J., & Tibbits, T.W. (1992). Importance of ‘blue’ photon levels for lettuce seedlings
grown under red-light-emitting diodes. HortScience, 27, 427-430.
http://www.ncbi.nlm.nih.gov/pubmed/11537611
Hu, C.M., & Hou, X.L. (2010). Relationship of major nutrient components with low temperature tolerance in
non-heading Chinese cabbage. Journal of Nanjing Agricultural University, 33, 37-41. (in Chinese with English
Abstract). http://nauxb.njau.edu.cn
Jaimez, R.E., & Rada, F. (2011). Gas exchange in Sweet Pepper (Capsicum chinense Jacq) under different light
conditions. Journal of Agricultural Science, 3, 134-142. http://dx.doi.org/10.5539/jas.v3n3p134
Kaneko-Ohashi, K., Fujiwara, K., Kimura, Y., Matsuda, R., & Kurata, K. (2004). Effects of red and blue LEDs
low light irradiation during low temperature storage on growth, ribulose-1, 5-bisphosphate carboxylase/oxygenase
content, chlorophyll content and carbohydrate content of grafted tomato plug seedlings. Environmental Control in
Biology, 42, 65-73. http://sciencelinks.jp/j-east/article/200704/000020070407A0032357.php
Kim, H.H., Goins, G.D., Wheeler, R.M., & Sager, J.C. (2004). Green-light supplementation for enhanced lettuce
growth under red- and blue-light-emitting diodes. HortScience, 39, 1617-1622.
http://www.ncbi.nlm.nih.gov/pubmed/15770792
Kim, H.H., Wheeler, R.M., Sager, J.C., Goins, G.D., & Norikane, J.H. (2006). Evaluation of lettuce growth using
supplemental green light with red and blue light-emitting diodes in a controlled environment-a review of
research at Kennedy Space Center. Acta Horticulturae, 711, 111-119.
http://www.actahort.org/members/showpdf?booknrarnr=711_11
Kolawole, O. M., Kayode, R. M. O., & Aina, J. (2010). The drying effect of varying light frequencies on the
proximate and microbial composition of tomato. Journal of Agricultural Science, 2, 214-224.
http://www.ccsenet.org/jas
Kowallik, W. (1982). Blue light effects on respiration. Annual Review of Plant Physiology, 33, 51-72.
http://dx.doi.org/10.1146/annurev.pp.33.060182.000411
Kurilcik, A., Miklusyte-Canova, R., Dapkuniene, S., Zilinskaite, S., Kurilcik, G., Tamulaitis, G., Duchovskis, P.,
& Zukauskas, A. (2008). In vitro culture of Chrysanthemum plantlets using light-emitting diodes. Central
European Journal Biology, 3, 161-167. http://dx.doi.org/10.2478/s11535-008-0006-9
Lefsrud, M.G., Kopsell, D.A., & Sams, C.E. (2008). Irradiance from distinct wavelength light-emitting diodes
affect secondary metabolites in Kale. HortScience, 43, 2243-2244. http://trace.tennessee.edu/utk_planpubs/6

268 ISSN 1916-9752 E-ISSN 1916-9760

www.ccsenet.org/jas Journal of Agricultural Science Vol. 4, No. 4; 2012

Li, H. M., Xu, Z. G., & Tang, C. M. (2010). Effect of light-emitting diodes on growth and morphogenesis of
upland cotton (Gossypium hirsutum L.) plantlets in vitro. Plant Cell Tissue Orgure Culture, 103, 155–163.
http://dx.doi.org/10.1007/s11240-010-9763-z
Li, Q., & Kubota, C. (2009). Effects of supplemental light quality on growth and phytochemicals of baby leaf
lettuce. Environmental and Experimental Botany, 67, 59-64. http://dx.doi.org/10.1016/j.envexpbot.2009.06.011
Lichtenthaler, H., & Wellburn, A. (1983). Determinations of total carotenoids and chlorophylls a and b of leaf
extracts in different solvents 603rd meeting, Liverpool England.
Liu, X.Y., Guo, S.R., Xu, Z.G., Jiao, X.L., & Takafumi, T. (2011). Regulation of chloroplast ultrastructure,
cross-section anatomy of leaves, and morphology of stomata of cherry tomato bydifferent light irradiations of
light-emitting diodes. Hortscience, 46, 1-5. http://hortsci.ashspublications.org/content/46/2/217.full
Massa, G.D., Kim, H.H., Wheeler, R.M., & Mitchell, C.A. (2008). Plant productivity in response to LED lighting.
HortScience, 43, 1951–1956. http://hortsci.ashspublications.org/content/43/7/1951.full
Menard, C., Dorais, M., Hovi, T., & Gosselin, A. (2006). Developmental and physiological responses of tomato
and cucumber to additional blue light. Acta Horticulturae, 711, 291-296.
http://www.actahort.org/books/711/711_39.htm
Mockler, T.C., Guo, H., Yang, H., Duong, H., & Lin, C. (1999). Antagonistic action of Arabidopsis cytochromes
and phytochrome B in the regulation of floral induction. Development, 126, 2073-2082.
http://dx.doi.org/10.1190/1.1438986
Morrow, R.C. (2008). LED Lighting in Horticulture. HortScience, 43, 1947-1950.
http://hortsci.ashspublications.org/content/43/7/1947.full
Okamoto, K., Yanagi, T., & Kondo, S. (1997). Growth and morphogenesis of lettuce seedlings raised under
different combinations of red and blue light. Acta Horticulturae, 435, 149-157.
http://www.actahort.org/books/435/435_14.htm
Poudel, P.R., Kataoka, I., & Mochioka, R. (2008). Effect of red-and blue-light-emitting diodes on growth and
morphogenesis of grapes. Plant Cell Tissue Organ Culture, 92, 147-153.
http://dx.doi.org/10.1007/s11240-007-9317-1
Saebo, A., Krekling, T., & Appelgren, M. (1995). Light quality affects photosynthesis and leaf anatomy of birch
plantlets in vitro. Plant Cell Tissue Organ Culture, 41, 177-185. http://dx.doi.org/10.1007/BF00051588
Schuerger, A.C., Brown, C.S., & Stryjewski, E.C. (1997). Anatomical features of pepper plants (Capsium
annuum L.) grown under red lightemitting diodes supplemented with blue or far-red light. Annals of Botany, 79,
273-282. http://dx.doi.org/10.1006/anbo.1996.0341
Senger, H. (1982). The effect of blue light on plants and microorganisms. Photochemistry and Photobiology, 35,
911-920. http://dx.doi.org/10.1111/j.1751-1097.1982.tb02668.x
Stutte, G.W. (2009). Light-emitting diodes for manipulating the phytochrome apparatus. HortScience, 44,
231–234. http://hortsci.ashspublications.org/content/44/2/231.full
Stutte, G.W., Edney, S., & Skerritt, T. (2009). Photoregulation of bioprotectant content of red leaf lettuce with
light-emitting diodes. HortScience, 44, 79-82. http://hortsci.ashspublications.org/content/44/1/79.full
Takahashi, K., Fujino, K., Kikuta, Y., & Koda, Y. (1995). Involvement of the accumulation of sucrose and the
synthesis of cell wall polysaccharides in the expansion of potato cells in response to jasmonic acid. Plant Science,
111, 11-18. http://dx.doi.org/10.1016/0168-9452(95)04222-G
Tamulaitis, G., Duchovskis, P., Bliznikas, Z., Breive, K., Ulinskaite, R., Brazaityte, A., Novickovas, A., &
Zukauskas, A. (2005). High-power light-emitting diode based facility for plant cultivation. Journal of Physics D:
Applied Physics, 38, 3182-3187. http://dx.doi.org/10.1088/0022-3727/38/17/S20
Tanaka, M., Takamura, T., Watanabe, H., Endo, M., Yanagi, T., & Okamoto, K. (1998). In vitro growth of
Cymbidium plantlets cultured under super bright and blue light-emitting diodes (LEDs). Journal of Horticultural
Science and Biotechnology, 73, 39-44. http://www.defra.gov.uk/
Yanagi, T., Okamoto, K., & Takita. S. (1996). Effects of blue, red and blue/red lights of two different PPF levels
on growth and morphogenesis of lettuce plants. Acta Horticulturae, 440, 117-122.
http://www.ncbi.nlm.nih.gov/pubmed/11541565
Yang, X.J., Liu, S.Q., Zhang, Z.K., Ma, L., Zhang, Y., & Wei, H. (2010). Effects of different light emitting

Published by Canadian Center of Science and Education 269

www.ccsenet.org/jas Journal of Agricultural Science Vol. 4, No. 4; 2012

diode sources on nutritional quality of garlic seedling. Acta Nutrimenta Sinica, 32, 518-520. (in Chinese with
English Abstract). http://mall.cnki.net/magazine/Article/YYXX201005031.htm
Yorio, N.C., Goins, G.D., Kagie, H.R., Wheeler, R.M., & Sager, J.C. (2001). Improving spinach, radish, and
lettuce growth under red light-emitting diodes (LEDs) with blue light supplementation. HortScience, 36,
380-383. http://www.ncbi.nlm.nih.gov/pubmed/12542027
Zhang, L.W., Liu, S.Q., Zhang, Z.K., Yang, R., & Yang, X.J. (2010). Dynamic effects of different light qualities
on pea sprouts quality. Northern Horticulture, 8: 4-7. (in Chinese with English Abstract).
http://www.cnki.com.cn/Article/CJFDTOTAL-BFYY201008003.htm
Zhang, Y. S., Huang, X., & Chen, Y. F. (2009). Experimental course of plant physiology. Higher Education Press,
Beijing (in Chinese).

Table 1. The effects of different lights on the mass of non-heading Chinese cabbage seedling growth under a
PPFD level of 80 μmol m–2s–1 for 60 days
Light Shoot fresh mass Shoot dry mass Root fresh mass Root dry mass
treatment (g) (g) (g) (g)
FL 1.38b 0.23c 0.13d 0.02d
B+R 5.28a 0.53b 0.95b 0.11b
B 6.67a 0.49b 0.61c 0.08c
R 5.93a 0.73a 1.08a 0.14a
Different letters within the column indicate significant differences at P<0.05 according to Tukey’s test.

Table 2. The effects of sunlight on the mass of non-heading Chinese cabbage seedlings that were first grown at a
PPFD level of 80 μmol m–2s–1 for 30 days and then transferred to sunlight for 30 days
Light Shoot fresh mass Shoot dry mass Root fresh mass Root dry mass
treatment (g) (g) (g) (g)
FL 3.69c 0.33c 0.33c 0.04d
B+R 10.95b 0.96a 0.93b 0.15a
B 11.79b 0.75b 0.90b 0.10b
R 13.60a 0.80b 1.40a 0.06c
Different letters within the column indicate significant differences at P<0.05 according to Tukey’s test.

Table 3. The effects of different lights on the mass of non-heading Chinese cabbage seedlings that were first
grown at a PPFD level of 80 μmol m–2s–1 (weak light) for 30 days, then under sunlight for 30 days and, finally,
under weak light at 61 days until flowering
Light Shoot fresh mass Shoot dry mass Root fresh mass Root dry mass
treatment (g) (g) (g) (g)
FL 0.61c 0.04c 0.04c 0.01c
B+R 6.83b 0.71b 1.36b 0.18b
B 9.73a 0.76ab 1.21b 0.20b
R 6.57b 0.87a 1.71a 0.24a
Different letters within the column indicate significant differences at P<0.05 according to Tukey’s test.

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Table 4. The Number of flower buds and open flowers per plant in non-heading Chinese cabbage seedlings grew
under different lights until flowering (90 -120 days)
Light Numbers of open Flowers Numbers of flower buds
treatment 0d 10d 20d 30 d 0d 10d 20d 30d
FL 0a 0b 5bc 8a 2a 4a 0b 0a
B+R 2a 5a 9ab 11a 3a 5a 2ab 2a
B 0a 0b 3c 9a 1a 7a 7a 2a
R 2a 5a 10a 11a 3a 5a 2ab 0a
Different letters within the column indicate significant differences at P<0.05 according to Tukey’s test.
2.5
a
2 a ab
bc
a FL
1.5 a c
Values

ab B+R
b B
1
a R
bc ab ab a
bc
0.5 c c
0
(mg/g) (mg/g) (mg/g) (mg/g)

Chl. a Chl. b Chl.(a+b) Car

2
1.8 a
a a
1.6 b a
1.4 a FL
a
1.2 a a
Values

B+R
1
B
0.8
a R
0.6 a a a a a a a
0.4
0.2
0
(mg/g) (mg/g) (mg/g) (mg/g)

Chl. a Chl. b Chl.(a+b) Car

2.5
ab a
2 c ab
FL
a a b
1.5
Values

a B+R
a B
1 ab a R
c a a
c a a
0.5

0
（mg/g） (mg/g) (mg/g) (mg/g)

Chl. a Chl. b Chl.(a+b) Car

Figure 1. The effects of different lights on the pigment concentrations of non-heading Chinese cabbage seedlings
with different PPFD levels, growth under LEDs and FL at a PPFD level of 80 μmol m–2s–1 for 60 days (a),
initial growth at a PPFD level of 80 μmol m–2s–1 for 30 days followed by sunlight for 30 days (b) and initial
growth at a PPFD level of 80 μmol m–2s–1 for 30 days followed by sunlight for 30 days and then weak light at 61
days until flowering (c)
Different letters within the column indicate significant differences at P<0.05 according to Tukey’s test.

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35
a
30
b
25 a
a a a
Vitamin C (mg/g)

20 b b
b
c
15 d c
c
10 b
d
5

0
FL B+R B R
Light treatment

Figure 2. The effects of different lights on the vitamin C concentration of non-heading Chinese cabbage
seedlings with different PPFD levels, growth under LEDs and FL at a PPFD level of 80 μmol m–2s–1 for 60
days (a), initial growth at a PPFD level of 80 μmol m–2s–1 for 30 days followed by sunlight for 30 days (b) and
initial growth at a PPFD level of 80 μmol m–2s–1 for 30 days followed by sunlight for 30 days and then weak light
at 61 days until flowering (c)
Different letters within the column indicate significant differences at P<0.05 according to Tukey’s test.

25

20 a
Souble protein (mg/g)

15 a
b
a a a a
10 a b
a a
c bc c
5 b

0
FL B+R B R
Light treatment

Figure 3. The effects of different lights on the soluble protein concentration of non-heading Chinese cabbage
seedlings with different PPFD levels, growth under LEDs and FL at a PPFD level of 80 μmol m–2s–1 for 60
days (a), initial growth at a PPFD level of 80 μmol m–2s–1 for 30 days followed by sunlight for 30 days (b) and
initial growth at a PPFD level of 80 μmol m–2s–1 for 30 days followed by sunlight for 30 days and then weak light
at 61 days until flowering (c)
Different letters within the column indicate significant differences at P<0.05 according to Tukey’s test.

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25
a
20 a
a

b FL
15 b
Values

B+R

c B
10 c c
R

a
5 b d
c
d

0
(mg/g) (mg/g) (mg/g)

Surcose Soluble Sugar Starch

120

100
b a

80 b FL
a a a
Values

B+R
60
b B
R
40

b c
20 a a a
b

0
(mg/g) (mg/g) (mg/g)

Surcose Soluble Sugar Starch

80 a

70 c
a
60 a

50 FL
b b
Values

B+R
40
B
30 R
a a
20 c c
c
b
10 c

0
(mg/g) (mg/g) (mg/g)

Surcose Soluble Sugar Starch

Figure 4. The effects of different lights on the photosynthesis production of non-heading Chinese cabbage
seedlings with different PPFD levels, growth under LEDs and FL at a PPFD level of 80 μmol m–2s–1 for 60
days (a), initial growth at a PPFD level of 80 μmol m–2s–1 for 30 days followed by sunlight for 30 days (b) and
initial growth at a PPFD level of 80 μmol m–2s–1 for 30 days followed by sunlight for 30 days and then weak light
at 61 days until flowering (c)
Different letters within the column indicate significant differences at P<0.05 according to Tukey’s test.

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