40

This is an open access article

under the CC BY license
Creative Commons

Light sources indoor benefit the growth and development of pepper

cultivars
Marcos Vinícius Marques Pinheiro1 , Maria Inês Diel2*, Oscar Valeriano Sanchéz Valera3, Leonardo Antonio Thiesen4,
Anderson Rafael Webler5, Guilherme Masarro-Araujo5, Diéssica Letícia Junges5, Tainara Gris6, Denise Schmidt5

10.1590/0034-737X202370030005

ABSTRACT
Light is a determining factor in plant morphophysiology, as it influences the growth and development of agricultural
crops. Thus, the objective was to investigate the effects of light sources on the growth and development of “biquinho”
pepper cultivars. The experiment was performed in two stages (Exp1 and Exp2), evaluating two cultivars (BRS Moema;
Airetama biquinho) in five light sources (white LEDs, red LEDs, blue LEDs, red/blue LEDs, and fluorescent lamps).
In Exp1 the plants were kept in a controlled condition, with a completely randomized design, in a 2x5 factorial scheme
(cultivars x light sources) until 76 days after emergence (DAE), in which growth variables and photosynthetic pigments
were evaluated. In Exp2, the plants were removed from the above conditions and transplanted in pots, being kept in
greenhouse for more 76 DAE. At 152 DAE, the same variables as Exp1 were evaluated, as well as gain of shoot fresh
and dry mass, and gain of root fresh and dry mass. In both experiments, growth variables were affected by cultivar and
the light sources, however, the behavior did not follow the same trend for all variables, indicating that the light quality
influences the growth of the crops, and impacting during greenhouse conditions.
Keywords: Capsicum chinense; light quality; light-emitting diodes (LEDs); photosynthetic pigments.

INTRODUCTION
The study of the physiology of plants and their behavior genesis and physiology, and the quality provided may be
in different environments allowed humans to both select able to influence plant productivity. The visible light range,
plants and modify their environment to obtain greater between 400 and 700 nm, is known as photosynthetically
productivity and other desired characteristics. However, active radiation, with the bands of red (600 to 700 nm) and
environmental factors, ie water availability, nutrients and blue (400 to 500 nm) being the ones that most affect pho-
the quality and quantity of light are important for plant tosynthesis, because the molecules of chlorophyll absorb
growth and development (Guo et al., 2016; Zhang et al., wavelengths in these bands, while the others are usually
2019). reflected or transmitted.
Among these environmental factors, light stands out Recent innovations in plant growth lighting, particular-
as an important regulator of plant development, morpho- ly through light-emitting diode (LED) technology, allow

_______________________________________________
Submitted on May 20th, 2022 and accepted on September 24th, 2022.
1 Universidade Estadual do Maranhão, São Luís, Maranhão, Brazil. [email protected]
2 Universidade Federal do Pampa, Itaqui, Rio Grande do Sul, Brazil. [email protected]
3 Instituto Nacional do México, Instituto Tecnológico Superior de Zongolica, Veracruz, Mexico. [email protected]
4 Universidade Federal de Santa Maria, Santa Maria, Rio Grande do Sul, Brazil. [email protected]
5 Universidade Federal de Santa Maria, Frederico Westphalen, Rio Grande do Sul, Brazil. [email protected]; [email protected]; [email protected]; denise@
ufsm.br
6 Universidade Federal de Santa Catarina, Florianópolis, Santa Catarina, Brazil. [email protected]
*Corresponding author: [email protected]

Rev. Ceres, Viçosa, v. 70, n. 3, p. 40-53, may/jun, 2023

 Light sources indoor benefit the growth and development of pepper cultivars 41

for the creation of customized light spectra, bringing 2016). The yellowish-green light (490-550 nm) is also
spectra into the decision-making process when a grower photosynthetically active radiation (Lin et al., 2020). In
or researcher is selecting agricultural lighting (Claypool & Dendranthema grandiflorum (chrysanthemum) under red
Lieth, 2020). LEDs have the flexibility to provide specific LEDs, increased in height and larger internodes, under
wavelengths with the possibility of selecting the emission blue/red LEDs there was largest fresh weight and greater
peak that most closely matches the absorption of photo- plant growth, and reduced plant growth kept under blue
receptors, and thus faster and more favorable results for LEDs (Kim et al., 2004). In Vaccinium corymbosum the
researchers and plant producers (Gupta & Jatothu, 2013; red light promoted greater rooting and elongation of the
Miler et al., 2019). Using light sources based on LEDs it is stem, and the combinations of blue/red LEDs benefited the
possible to regulate and control the physiological aspects accumulation of plant biomass in vitro (Hung et al., 2016).
of plant growth, such as photosynthesis and/or photomor- The use of far red inside the canopy of tomato plants
phogenesis (Gupta & Jatothu, 2013). resulted in stem elongation, difference in leaf morphology,
In addition, the light emitted from the LEDs is an abiotic greater length/width ratio of the leaves, largest leaf area
elicitor capable of being used in plant morphogenesis, as it and increase of 7 to 12% in the production of ripe fruits
has high specificity in wavelengths, allowing researchers (Zhang et al., 2019). In peppers, the effects of LED lights
to eliminate other wavelengths found in white light, and on growth, yield and fruit quality were tested, with an
thus study the stimulating them in the architecture of the increase in fruit production, in the dry matter content of
plant and in the productive responses. The use of light the fruits and in the content of fruit-promoting compounds
wavelengths promotes different morphogenetic and pho- in the fruits (i.e. total phenolic content, carotenoid content
tosynthetic responses that may vary among plant species and antioxidant activities) (Guo et al., 2016).
(De Hsie et al., 2019), and in terms of plant production, In Capsicum annuum, plant biomass was reduced
there are only a few applications where a spectrum of pure when the plants remained under red LEDs in the absence
monochrome light produces good growth results, such as of the blue wavelength, producing characteristics such as
keeping plants compact or retarding plant growth (Miler greater height and greater mass of the stem, suggesting
et al., 2019). that the plants require supplementary radiation especially
LED lighting technologies for plant cultivation are in the blue region for the normal growth and development
rapidly evolving, and lamps for indoor cultivation are (Brown et al., 1995). Even with these results, the same
often designed to optimize their light emissions in the authors confirm the physiological and morphological
photosynthetically active spectrum (i.e. red and blue), effects of the quality of light, and they can vary according
to reduce energetic requirements for satisfactory yield to the species. For cultivars of biquinho pepper (Capsicum
(Pennisi et al., 2020). This technology has been widely chinense), light spectra were studied during the seed ger-
used in horticultural facilities in recent years, however, mination process, concluding that the luminous environ-
the influence of light quality should be further explored ment in green, red and far red colors positively influence
(Yang et al., 2018). Red and blue lights are currently two the germination and the amount of abnormal plants was
types of light spectra that have been most studied on plant increased in the dark (Diel et al., 2019). For this species,
photobiology (Zhang et al., 2020). this is the only report, and it is crucial to study the influ-
Plants respond differently to each photosynthetic ence of light source on the morphogenesis and physiology
pigment, the quality of light has varying influences on of this culture.
plant growth and development. The red light (660 nm) and Several studies have revealed the effects of light on
the distant red light (730 nm) affect phytochrome, which the biomass of different species, using LEDs of specific
affects from seed germination to flowering, fruiting and wavelengths. For this, the question remains: which band
aging (Oren et al., 2021). The blue-violet light (400-490 of the light spectrum can influence the morphogenesis of
nm) regulates plant growth through cryptochromes and Capsicum chinense? The aim of the study was to evaluate
phototropins, affecting the growth and development of the effects of light sources indoor on the morphogenesis
the root system, and plant stem growth (Pedmale et al., (growth and development) of two cultivars of pepper.

Rev. Ceres, Viçosa, v. 70, n. 3, p. 40-53, may/jun, 2023

42 Marcos Vinícius Marques Pinheiro et al.

MATERIALS AND METHODS nitrate and 15 g 1000 L-1 of iron EDDHA, Yara®), and
electrical conductivity of the solution maintained at 600
2.1 Study area and growth conditions
µS. The plants remained in growth room conditions until
The experiment was carried out at the Federal University the beginning of flowering, which was determined when
of Santa Maria UFSM campus Frederico Westphalen - Rio 50% of the plants in the plot emitted flowers, that is, with
Grande do Sul, Brazil. The geographical location of this 76 DAE.
region is 27º22’S, 53º25’O at 480 m of altitude. According
to the Köppen classification, the region’s climate is Cfa, 2.2.2 Experimental design and variables analyzed
humid subtropical (Alvares et al., 2013). The experimental design used was entirely random in a
For the conduct of the experiment, two steps were 2 x 5 factorial scheme, with two cultivars of pepper (BRS
performed (Experiment I and II), the first was conducted in Moema and Airetama Biquinho) and five light sources
a growth room with controlled temperature, humidity and (fluorescent lamps, white LEDs, red LEDs, blue LEDs, and
light intensity conditions and the second in a greenhouse. red/blue LEDs), totaling ten treatments with four repeti-
tions per treatment, the experimental unit being composed
2.2 Experiment I - Does the light source indoor
of four plants, totaling 16 plants per treatment.
benefit the vegetative process of biquinho pepper
At 76 DAE and before transplantation (Experiment II)
cultivars?
were evaluated: plant height (PH, cm), number of leaves
2.2.1 Plant material and propagation (NL), shoot fresh mass (SFM, g), shoot dry mass (SDM, g),
root fresh mass (RFM, g), root dry mass (RDM, g), root
Seeds of two cultivars of “biquinho” pepper, Capsicum
volume (RV, mL), root length (RL, cm), leaf area (LA, m²)
chinense, (BRS Moema and Airetama Biquinho) were
and photosynthetic pigments [chlorophyll a (Chla), chloro-
used, characterized by red and yellow fruits when ripe,
phyll b (Chlb), total chlorophylls (ChlT), carotenoids (Car)
respectively. The seeds were pre-germinated on Germitex®
and total chlorophyll/carotenoid ratio (ChlT/Car)].
paper in transparent plastic Gerbox® and kept in a growth
Before transplanting the plants to the pots, half of them
chamber (BOD type) at a constant temperature of 25 °C, 12
were separated for destructive evaluations of Experiment I
hours of light and 12 hours of photoperiod with 36 µmol
and the other half for Experiment II, being evaluated at the
m-2 s-1 irradiance from two fluorescent lamps (“Luz do
end of 152 DAE.
Dia Especial”, 40 W, Osram®, Brazil). To break the seed
The number of leaves and plant height were evaluated
dormancy, KNO3 solution (2%) was added, following the
from the emission of the first expanded leaf to the first fork
standards of MAPA (2009), and after about 15 days after
in the main stem of the plant. The root volume was calcu-
emergence (DAE), when 50% of the seeds showed root
lated immediately after calculating the fresh mass, from the
protrusion, they were transplanted into transparent plastic
displacement of water in a graduated cylinder after submer-
cups (300 mL) filled with Carolina® commercial substrate.
sion of the root system. The root length was evaluated by
From the transplant to plastic cups, the seedlings
measuring the length of the main root (pivoting root).
were kept in a growth room, with a temperature of 25 ±
The leaf area (LA, cm²) was determined using an LA
2 ºC, a photoperiod of 16 hours of light and eight hours
integrator [model Licor LI- 3100 (LI-COR®, Lincoln, Ne,
of darkness, with a luminous intensity of 72 µmol m-2 s-1
USA)]. For this, all the leaves were separated from the stem
from different light sources: four fluorescent lamps (“Luz
of the plants, being passed one by one in the analyzer.
do Dia Especial”, 40 W, Osram®, Brazil); two tube lamps
(TECNAL TECLAMP®, Piracicaba, Brazil) of different 2.3 Experiment II - Mass gains and quality of
colors [white LEDs, red LEDs (660 nm), blue LEDs (450 plants subjected to light source during the vegeta-
nm), red/blue LEDs (660 and 450 nm, with a ratio of 60 tive process of cultivars of biquinho pepper
and 40%, respectively).
2.3.1 Plant material
The plants were irrigated manually keeping the humid-
ity constant (to avoid water stress), and the plant nutrition At 76 DAE the plants were removed from the growth
was carried out via fertigation, with nutrient solution (200 room conditions and transplanted in black pots (5 L) filled
g 1000 L-1 of HidrogoodFert®, 250 g 1000 L-1 of calcium with typical dystrophic Red Latosol soil, and kept in a

Rev. Ceres, Viçosa, v. 70, n. 3, p. 40-53, may/jun, 2023

 Light sources indoor benefit the growth and development of pepper cultivars 43

greenhouse with a metallic structure in the shape of an the SPERCORD 50/Plus® spectrophotometer (Analytik-
arch, 3.5 m high, 10 and 20 m wide and long, covered with jena, Germany) using a 10 mm quartz cuvette. Wellburn
plastic film 150 microns thick, remaining for another 76 (1994) methodology was used to establish wavelength and
days, with a total of 152 DAE (fruit ripening; 50% of plants equations to calculate the concentrations of chlorophylls
with ripe fruit). [a, b, total (a + b)] and carotenoids and the unit used was
The plants were irrigated manually, always maintaining in µg cm-2.
constant humidity to avoid water stress. For this purpose,
2.4 Statistical analysis
the same fertigation solution mentioned above was used,
remaining until 152 DAE. The variables of both experiments (I and II) were
subjected to analysis of variance and the treatment means,
2.3.2 Experimental design and variables analyzed
when significant, were separated by the Tukey test at 5%
This experiment was conducted in a randomized block probability of erro. In addition, based on the desired values
design in a 2 x 5 factorial scheme, with two cultivars of of the variables studied, the MGIDI index was used to select
biquinho pepper (BRS Moema and Airetama Biquinho) the best treatments (Olivoto & Nardino, 2020) and were
[∑ j =1(γ ij − γ j ) 2 ]0.5 , where
f
and plants from five light sources (cited in Experiment I), calculated as follows: MGIDI
= i

totaling ten treatments with ten repetitions per treatment, MGIDIi is the multi-trait genotype-ideotype distance index
and each repetition composed of a plant. for the ith treatment; γij is the score of the ith treatment in
At 152 DAE, were evaluated: plant height (PH, cm), the j th factor (i = 1; 2; :::; t; j = 1; 2; :::; f), being t and f
number of leaves (NL), shoot fresh mass (SFM, g), shoot the number of treatments and factors, respectively; and γj
dry mass (SDM, g ), root fresh mass (RFM, g), root dry is the j th score. Treatments that have the lowest calculated
mass (RDM, g), root volume (RV, mL), root length (RL, indices are the best. Statistical analyzes were performed
cm), photosynthetic pigments [chlorophyll a (Chla), chlo- with the aid of packages metan (Olivoto & Lúcio, 2020),
rophyll b (Chlb), total chlorophylls (ChlT), carotenoids ggplot2 (Wickham, 2016), ExpDes (Ferreira et al., 2021)
(Car) and total chlorophyll/carotenoid ratio (ChlT/Car)] available in the R program (R Development Core Team,
and fresh fruit mass (FFM, g) and dry fruit mass (DFM, 2021).
g). The mass gain was calculated as a percentage for SFM,
SDM, RFM, SDM shoot fresh mass (SFM, g), shot dry RESULTS
mass (SDM, g), root fresh mass (RFM, g) and shot dry
3.1 Experiment I - Light source indoor benefit the
mass (SDM, g), at 76 DAE and 152 DAE (Experiment I
vegetative process of biquinho pepper cultivars
and II). That is, during the period between and greenhouse
conditions (76 DAE and 152 DAE, respectively), the mass- From the analysis of variance, there was a significant
es were related, being calculated as a percentage: gain of interaction between cultivars x light sources for the
shoot fresh mass (%, GSFM), gain of shoot dry mass (%, variables PH, NL, SFM, RFM, SDM, RDM and LA.
GSDM), gain of root fresh mass (%, GRFM), gain of root The variables RL and RV were significant only for the
dry mass (%, GRDM). cultivars factor, and CR only for light sources. For pho-
tosynthetic pigments, there was significant interaction
2.3.3 Photosynthetic pigments
between cultivars x light sources for Chla, Chlb, ChlT,
To determine photosynthetic pigments (Experi- Car, ChlT/Car.
ment I and II), chlorophyll a, chlorophyll b, total chloro- The growth variables were affected by both the cultivar
phyll and carotenoids, five disks of 5 mm diameter each and the light sources, in the first stage of the experiment
were obtained from the second green leaf (from the apex to (Figure 1). For the PH variable, the cultivar BRS Moema
the base) and incubated in glass test tubes containing 5 mL was superior under blue (15.5 cm) and red LEDs (11.37
of dimethylsulfoxide (DMSO) saturated with calcium car- cm) but obtained lower averages than Airetama in the flu-
bonate (CaCO3), kept in the dark at room temperature for orescent light source (8.21 cm) (Figure 1A). The same pat-
48 hours, following the modified methodology of Santos tern occurred for the variables SFM and SDM (Figure 1C,
et al. (2008), totaling four repetitions per treatment. After 1D , respectively). BRS Moema, when kept under LEDs,
48 hours of darkness, absorbance was determined using also obtained higher NL values except for the fluorescent

Rev. Ceres, Viçosa, v. 70, n. 3, p. 40-53, may/jun, 2023

44 Marcos Vinícius Marques Pinheiro et al.

light source, where NL was higher for the cultivar Airetama LEDs (Figure 1E).
(Figure 1B). Differently, RDM presented another behavior For the variable RL, the cultivar Airetama had a longer
when compared to the other variables, with higher averages root length when grown under white LEDs (Figure 2A),
in the cv. Airetama when grown under Blue LEDs (Figure and shorter when the plants remained under fluorescent
1F). For the RFM variable, the inferior performance was and red LEDs (Figure 2C), the same being observed for the
observed for the BRS Moema cultivar grown under white variable RV (Figure 2B).

Figure 1: Means of growth variables of biquinho pepper cultivars under different light sources in experiment I: A) plant height (PH,
cm), B) number of leaves (NL), C) shoot fresh mass (SFM, g), D) shoot dry mass (RFM, g), E) root fresh mass (SDM, g), F) root dry
mass (RDM, g). Lower case letters differ in light sources and upper-case letters differ in cultivars by Tukey’s test, at 5% significance.

The cultivar BRS Moema showed superior results of light sources blue/red LEDs and white LEDs (Figure 3A).
LA when cultivated under blue LEDs and red LEDs. The For chlorophyll a (Chla), the cultivar BRS Moema obtained
cultivar Airetama performed better under fluorescent light, superior results when cultivated under blue LEDs and blue/
while both cultivars showed low performance under the red LEDs, and inferior performance under fluorescent light,

Rev. Ceres, Viçosa, v. 70, n. 3, p. 40-53, may/jun, 2023

 Light sources indoor benefit the growth and development of pepper cultivars 45

in which the cultivar Airetama obtained superior results comparing with all the other light sources studied. BRS
(Figure 3B). BRS Moema presented superior results of Moema had a similar performance when grown under
chlorophyll b (Chlb) under blue LEDs when compared to blue, red and blue/red LEDs (Figure 3E). The ChlT/Car
Airetama, in the other light sources studied, the two cul- ratio showed significant differences between cultivars
tivars did not present significant differences. The cultivar only under blue LEDs, while within each BRS Moema
Airetama did not differ in any of the light sources, while cultivar it presented a higher ChlT/Car ratio under blue
the cultivar BRS Moema obtained a higher Chlb content LEDs, and Airetama did not show significant differences
in the blue light source (Figure 3C). Carotenoids showed between light sources (Figure 3F).
significant differences between cultivars in the blue/red By using the multi-trait index (MGIDI), we selected
LED light source, in which BRS Moema was superior to the 2 best treatments (20%) being T3 (Airetama biquinho
Airetama, and when kept under fluorescent light, the Aire- and fluorescent light) and T6 (BRS Moema and Blue
tama cultivar was superior to BRS Moema. It was also LEDs), showing the difference in the response of the
observed in the light sources in each cultivar, with BRS cultivars in relation to the type of light to which it was
Moema being superior under blue/red LEDs and Airetama submitted (Figure 7A). The two treatments were selected
under fluorescent (Figure 3D). with 90% of the variables with desired values, and only
For the variable ChlT, among the cultivars, BRS the RV variable did not have the desired selection (SD =
Moema was significantly superior to Airetama in the -2,35%). Thus, we can say that treatments T3 (Airetama
blue and blue/red LED light sources, while the cultivar biquinho and fluorescent light) and T6 (BRS Moema
Airetama was superior to BRS Moema in the fluorescent and blue LEDs) had a good performance in all analyzed
light source, in which it had superior performance when variables, except for RV.

Figure 2: Growth variables (A, C) root length (RL, cm) and (B) root volume (RV, mL) as a function of biquinho pepper cultivars under
different light sources in growing room. Lower case letters differ by Tukey’s test, at 5% significance.

Rev. Ceres, Viçosa, v. 70, n. 3, p. 40-53, may/jun, 2023

46 Marcos Vinícius Marques Pinheiro et al.

Figure 3: (A) leaf area (LA, m²), (B) chlorophyll a (Chla, µm cm-2), (C) chlorophyll b (Chlb, µm cm-2), (D) carotenoids (CAR, µm
cm-2), (E) total chlorophyll content (ChlT, µm cm-2), and F) total chlorophyll/carotenoid ratio (ChlT/Car) of biquinho pepper cultivars
grown under different light sources in growing room. Lower case letters differ in light sources and upper-case letters differ in cultivars
according to the Tukey's test, at 5% significance.

3.2 Experiment II - Mass gains and quality of significant effect. For the variables of mass gain (GSFM,
plants subjected to light sources during the vegeta- GSDM, GRFM, GRDM) there was a significant interaction
tive process of cultivars of biquinho pepper between cultivar x light sources.
The variable RV, after transplanting the seedlings to
From the analysis of variance, it was possible to observe pots kept inside the greenhouse, the behavior of the cultivar
that there was interaction between cultivars x light sources Airetama was superior to BRS Moema when grown under
for the growth variables RV, SFM, RSM, RDM. For PH, blue LEDs and red LEDs. In the other light sources, the
NL, SDM and RL there was no significant difference. For cultivars did not differ significantly. For each cultivar, Aire-
photosynthetic pigments, there was interaction between tama showed superior results when kept under blue LEDs
cultivars x light sources for the variables Chla, ChlT and and red LEDs, and the BRS Moema cultivar showed no
Car, whereas for Chlb and ChlT/Car ratio there was no significant differences for light sources (Figure 4A).

Rev. Ceres, Viçosa, v. 70, n. 3, p. 40-53, may/jun, 2023

 Light sources indoor benefit the growth and development of pepper cultivars 47

Figure 4: (A) root volume (RV, ml), (B) shoot fress mass (SFM, g), (C) root fresh mass (g), (RFM, D) root dry mass (RDM, g), E) fresh
fruit mass (FFM, g) and F) dry fruit mass (DFM, g).of piquet pepper cultivars at 152 DAE in greenhouse. Lower case letters differ in
light sources and upper-case letters differ in cultivars according to the Tukey test, at 5% significance.

As for photosynthetic pigments, contrary to what red LEDs. Within each cultivar, BRS Moema showed
was observed in Experiment I, lower levels of Chla were higher ChlT content when the plants were under red
observed for the BRS Moema cultivar when comparing and blue/red LEDs. Under blue LEDs, Airetama showed
Airetama under blue LEDs, however the fluorescent light superior performance (Figure 5B). There were significant
continued to be the one with the lowest Chla values of differences between cultivars for carotenoids, that is,
the BRS Moema cultivar (Figure 5A). For ChlT, among the cultivar Airetama when grown under blue LEDs and
cultivars, different from experiment I, Airetama was fluorescent lamps was superior to BRS Moema, while the
significantly superior to BRS Moema in blue LEDs and latter was significantly superior under red LEDs. Within
under fluorescent lamps, while the cultivar BRS Moema each cultivar, this variable followed the same trend found
was significantly superior to Airetama when kept under for the ChlT variable (Figure 5C). Analyzing the results

Rev. Ceres, Viçosa, v. 70, n. 3, p. 40-53, may/jun, 2023

48 Marcos Vinícius Marques Pinheiro et al.

obtained, we can infer that each cultivar has different trolled conditions (Experiment I) and the impact of these
characteristics regarding the production of photosynthetic on the growth and development of plants in greenhouse
pigments and in relation to each light source. cultivation (Experiment II) to infer about the increment and
The mass gain was significant for all treatments (light gain of the plants in those conditions.
sources x cultivar), for 76 days passed from the initial to For most of the variables analyzed, there was an
the final evaluation. For GSFM, the lowest mass gain of interaction between cultivars x light sources evaluated in
the cultivar Airetama was observed in the fluorescent light the study. It is known that pepper genotypes can respond
source (101.4%), while for the cultivar BRS Moema the differently to environmental stimuli, and depending on
lowest mass gains were observed under blue LEDs (57.3%) the environment, they can produce, for example, different
(Figure 6A). For GSFM, the cultivar Airetama had no dif- amounts of capsaicinoids and have differences in develop-
ferences between the light sources, while for the cultivar ment (Jeeatid et al., 2018).
BRS Moema higher values were observed when they were The cultivars showed different responses to the light
under fluorescent and white LEDs (1338.3 and 1240.1% sources in which they were submitted. For example, the
respectively). Another important factor was observed for plant height (PH) of the cultivar BRS Moema was higher
the fluorescent source, which showed low GSDM for the when they were associated with blue LEDs and red LEDs.
cultivar Airetama (332.6%), while for BRS Moema it was The spectrum of blue and red light also impacted the height
superior (Figure 6B). of petunia flowers, due to the elongation of the stem, due
For GRFM, the cultivar Airetama did not differ between to the blue light stimulating the production of gibberellins,
the light sources evaluated, while for the cultivar BRS and thus causing changes in height (Fukuda et al., 2016).
Moema, when they were subjected to fluorescent sources, it However, the wavelengths in blue and red can positively or
presented the greatest gains compared to LEDs (2578.3%). negatively affect the growth and development of the spe-
In comparison to cultivars within each light source, the cies, and this is because these wavelengths are responsible
differences were also related to gains in cultivation under for the activation of different genetic expressions in plants
fluorescent (Figure 6C). The same trend was observed (Jeeatid et al., 2018).
for the GRDM variable, in which the greatest gains were For growth variables, plants from cultivar BRS Moema
achieved under fluorescent lamps for the cultivar BRS showed superior results when grown under monocromatics
Moema (5213.6%) (Figure 6D). blue or red LEDs. In contrast, the cultivar Airetama showed
Regarding the multi-trait index (MGIDI), as in experi- greater growth when cultivated under a source of fluores-
ment I, the 2 best treatments were also selected (20%) being cent light and white LEDs; this reveals that the responses
T1 (Airetama biquinho and blue LEDs) and T3 (Airetama between genotypes of the same species may be different
biquinho and fluorescent light), showing the difference in (Jeeatid et al., 2018), and also demonstrated by MGIDI in
the response of the light used (Figure 7B). The two treat- Experiment I. Fluorescent lamps commonly applied, gen-
ments were selected with 60% of the variables with desired erally lack a wavelength in red, which is very important for
values, and the variables RL, NL, SFM and SDM did not the development of plants, for example in stem elongation
have the desired selection (SD = -5.3%, -9.57%, -2.37%, and phytochrome activity, while these lamps emit a lot in
and -0.56%, respectively). Thus, we can say that treatments the green and yellow light spectra, which are less efficient
T1 (Airetama biquinho and blue LEDs) and T3 (Airetama for plants (Miler et al., 2019). In Capsicum annuum, a
biquinho and fluorescent light) had a good performance in broad spectrum including red, green, and blue wavelengths
most of the variables analyzed. resulted in the highest shoot dry weight and plant compact-
ness, and under monochromatic blue LEDs, these authors
DISCUSSION observed a reduced leaf area under blue LEDs (Claypool &
This is the first report on the use of light source in the Lieth, 2020); differently from the present work, suggesting
cultivation of Capsicum chinense cultivars kept under con- that this may be a species-specific response.

Rev. Ceres, Viçosa, v. 70, n. 3, p. 40-53, may/jun, 2023

 Light sources indoor benefit the growth and development of pepper cultivars 49

Figure 5: (A) chlorophyll a (Chla, µm cm-2), B) total chlorophyll (ChlT, µm cm-2), C) carotenoids (Car, µm cm-2) from biquinho pepper
cultivars grown at 152 DAE in greenhouse. Lower case letters differ in light sources and upper-case letters differ in cultivars according
to the Tukey's test, at 5% significance.

Rev. Ceres, Viçosa, v. 70, n. 3, p. 40-53, may/jun, 2023

50 Marcos Vinícius Marques Pinheiro et al.

Figure 6: Mass gain as a percentage of 76 DAE to 152 DAE for (A) gain of shoot fresh mass (GSFM, %), (B) gain of shoot dry mass
(GSDM, %), (C) gain of root fresh mass (GRFM, %) and (D) gain of root dry mass (GRDM, %) for Airetama and BRS Moema cultivars
at 152 DAE in greenhouse. Lower case letters differ in light sources and upper-case letters differ in cultivars according to the Tukey's
test, at 5% significance.

Figure 7: Treatments ranking in ascending order for the MGIDI index. The selected treatments are shown in red color and the red
circle represents the cutpoint according to the selection pressure. T1: Airetama biquinho and Blue LEDs, T3: Airetama biquinho and
fluorescent light and T6: BRS Moema and Blue LEDs.

Rev. Ceres, Viçosa, v. 70, n. 3, p. 40-53, may/jun, 2023

 Light sources indoor benefit the growth and development of pepper cultivars 51

Chlorophylls are photosynthetic pigments that The increase gains attributed to the fluorescent and white
absorb light to provide photosynthetic energy to plants LEDs in the period; may be related to the phase in which
(Fang et al., 2022) and contribute to plant vigor due to the plant is at the moment (Chen et al., 2014). In contrast,
photosynthetic efficiency (Huang et al., 2013), mainly Ajdanian et al. (2019), observed superiority with the ap-
comprise chlorophyll a (Chl a), chlorophyll b (Chl b) and plication of blue and red light, as there was an increase in
carotenoids (Car). Chlorophyll (Chl) absorb energy from fresh and dry masses of cress (Lepidium sativum), as well
sunlight and convert the light into chemical energy (Björn, as their biomass compared to plants grown under natural
2015), while carotenoids have the function of collecting sunlight conditions.
light in the range of 450-570 nm and protecting plants Due to the large number of treatments tested (2 culti-
against photooxidative damage (Frank & Cogdell, 1996). vars and 5 light sources = 10 treatments) it can be difficult
For photosynthetic pigments, the cultivars showed different to select the treatments that present the greatest selection
responses, while the contents of Chla, Chlb and carotenoids differential, that is, desired characteristics. Using MGIDI
in the cultivar BRS Moema were higher results for the we have the possibility to find the best treatments in a
blue, red LEDs and their combination. Shin et al. (2008), simple way and based on a multitrait framework (Olivoto
also observed that pigment biosynthesis was increased in & Nardino, 2020).
Doritaenopsis plants grown with the plus of red and blue Light is an important signal element that regulates
LEDs. These same authors report that the use of this mixed plant morphology, physiology and development during
red and blue radiation can benefit the quality production the plant growth cycle (Chen et al., 2014), and the
of plants. The cultivar Airetama showed higher values application of different light sources and their effects on
of Chla and Chlb for the fluorescent light, in contrast on plants is positive, as they result in the improvement of
lettuce (Chen et al., 2014) and Doritaenopsis (Shin et al., the biochemical characteristics and stimulates the use of
2008) both under fluorescent lamps. It is therefore difficult these light sources in an effort to increase the quality of
to indicate a single spectrum of light suitable for the prop- the plant and understand biochemically what happens
agation of all plant species (Miler et al., 2019), and in the with this application, and how it influences plant mor-
present work, this response was species-specific in many of phology. And this is only possible due to advances in
the analyzed variables. technology, in which they must be constantly updated,
There was a change in the behavior of BRS Moema and simultaneously stimulates the use of these in new
plants at 76 DAE, when they were compared to 152 DAE studies and applications by researchers worldwide, and
for the variables SFM, RFM e SDM. At 76 DAE, these can also be used as allies in future plant production.
plants showed superiority when grown under blue LEDs,
however, at 152 DAE, this result changed, with those grown CONCLUSIONS
under fluorescent and white LEDs with greater increment; The results of the present study indicate that the light
suggesting that plant growth responses to light quality may sources emitted in part of the growth of the plants becomes
not only be associated with species or cultivars, but also the crucial during greenhouse conditions when these plants
plant growth stages (Chen et al., 2014). were transferred to soil. The treatments selected under
The use of artificial lighting has a positive effect on the greenhouse conditions are T3 (Airetama biquinho and flu-
growth and development of the biquinho pepper cultivars, orescent light) and T6 (BRS Moema and Blue LEDs) and
proven by the gain of differential biomass between them. in field conditions T1(Airetama biquinho and Blue LEDs)
The greatest gains of SFM and SDM for BRS Moema and T3 (Airetama biquinho and fluorescent light). Our
cultivar were under fluorescent and white LEDs, for the discovery achieved the proposed objective, demonstrating
variables RFM and RDM the greatest gains were observed that the use of LEDs is effective in the development and
in the fluorescent. The cultivar Airetama showed no differ- more vigorous growth of Capsicum chinense, and that this
ences (p > 0.05) for mass gain in different light sources. response is dependent cultivar.

Rev. Ceres, Viçosa, v. 70, n. 3, p. 40-53, may/jun, 2023

52 Marcos Vinícius Marques Pinheiro et al.

ACKNOWLEDGMENTS FINANCIAL SUP- Huang J, Qin F, Zang G, Kang Z, Zou H & Hu F (2013) Mutation of
OsDET1 increases chlorophyll content in rice. Plant Science, 210:241-
PORT AND FULL DISCLOSURE 249.
We thank the National Council for Scientific and Hung CD, Hong CH, Kimm SK, Lee KH, Park JY, Nam MW, Choi DH
& Lee HI (2016) LED light for in vitro and ex vitro efficient growth of
Technological Development and Coordination for the Im- economically important highbush blueberry (Vaccinium corymbosum
provement of Higher Education Personnel for fellowships L.). Acta Physiologiae Plantarum, 38:152.
Jeeatid N, Suriharn B, Techawongstien S, Chanthai S, Bosland PW &
and grants to the authors. Techawongstien S (2018) Evaluation of the effect of genotype-by-en-
vironment interaction on capsaicinoid production in hot pepper
CONFLICT OF INTERESTS hybrids (Capsicum chinense Jacq.) under controlled environment.
Scientia Horticulturae, 235:334-339.
There is no conflict of interests in carrying this research Kim SJ, Hahn EJ, Heo JW & Paek KY (2004) Effects of LEDs on net
photosynthetic rate, growth and leaf stomata of chrysanthemum plant-
and publishing this manuscript. lets in vitro. Scientia Horticulturae, 101:143-151.
Lin KH, Shih FC, Huang MY & Weng JH (2020) Physiological char-
REFERENCES acteristics of photosynthesis in yellow-green, green and dark-green
Chinese Kale (Brassica oleracea L. var. alboglabra Musil.) under
Ajdanian L, Babaei M & Aroiee H (2019) The growth and development
varying light intensities. Plants, 9:960.
of cress (Lepidium sativum) affected by blue and red light. Heliyon,
5:e02109. MAPA - Ministério da Agricultura, Pecuária e Abastecimento (2009)
Regras para análise de sementes. Available at: <https://www.gov.br/
Alvares CA, Stape JL, Sentelhas PC, Moraes Golçalves JL & Sparovek
agricultura/pt-br/assuntos/laboratorios/arquivos-publicacoes-labora-
G (2013) Koppen’s climate classification map for Brazil. Meteorolo-
torio/regras-para-analise-de-sementes.pdf/view>. Accessed on: March
gische Zeitschrift, 22:711-728.
15th, 2022.
Björn LO (2015) The evolution of photosynthesis and its environmental
Miler N, Kulus D, Woźny A, Rymarz D, Hajzer M, Wierzbowski K &
impact. In: Björn LO (Ed.) Photobiology. New York, Springer. p.207-
Nelke R Szeffs L (2019) Application of wide-spectrum light-emitting
230.
diodes in micropropagation of popular ornamental plant species: a
Brown CS, Schuerger AC & Sager JC (1995) Growth and photomor- study on plant quality and cost reduction. In Vitro Cellular & Develop-
phogenesis of pepper plants under red light-emitting diodes with mental Biology - Plant, 55:99-108.
supplemental blue or far-red lighting. Journal of the American Society
Mitchell CA & Sheibani F (2020) LED advancements for plant-factory
for Horticultural Science, 120:808-813.
artificial lighting. In: Kozai T, Niu G &¨Takagaki M (Eds.) Plant
Chen XL, Guo WZ, Xue XZ, Wang LC & Qiao XJ (2014) Growth and Factory. 2º ed. Amsterdam, Academic Press. p.167-184.
quality responses of “Green Oak Leaf” lettuce as affected by mono-
Olivoto T & Lúcio AD (2020) Metan: an R package for multi-environ-
chromic or mixed radiation provided by fluorescent lamp (FL) and
ment trial analysis. Methods in Ecology and Evolution, 11:783-789.
light-emitting diode (LED). Scientia Horticulturae, 172:168-175.
Olivoto T & Nardino M (2020) MGIDI: toward an effective multivariate
Claypool NB & Lieth JH (2020) Physiological responses of pepper
selection in biological experiments. Bioinformatics, 37:1383-1389.
seedlings to various ratios of blue, green, and red light using LED
lamps. Scientia Horticulturae, 268:109371. Oren SN, Timm M, Frank O, Mantovani O & Murik M (2021) Red/
far-red light signals regulate the activity of the carbon-concentrating
De Hsie BS, Bueno AIS, Bertolucci SKV, De Carvalho AA, Da Cunha
mechanism in cyanobacteria. Science Advances, 7:435.
SHB, Martins ER & Pinto JEBP (2019) Study of the influence of wave-
lengths and intensities of LEDs on the growth, photosynthetic pigment, Pedmale UV, Shan S, Huang C, Zander M, Cole BJ, Hetzel J, Ljung
and volatile compounds production of Lippia rotundifolia Cham in K, Reis ABP, Sridevi P, Nito K, Nery RJ, Ecker RJ & Chory J (2016)
vitro. Journal of Photochemistry and Photobiology, 198:111577. Cryptochromes interact directly with PIFs to control plant growth in
limiting blue light. Cell, 164:233-245.
Diel MI, Valera OVS, Pinheiro MVM, Thiesen LA, Meira D, Melo PJ,
Junges DL, Caron BO & Schmidt D (2019) Temperature and light Pennisi G, Pistillo A, Orsini F, Cellini A, Spinelli F, Nicola S, Fernandez
quality influence seed germination of two biquinho pepper cultivars JA, Crepaldi A, Gianquinto G & Marcelis LFM (2020) Optimal light
Bulgarian. Journal of Agricultural Science, 25:1007-1014. intensity for sustainable water and energy use in indoor cultivation
of lettuce and basil under red and blue LEDs. Scientia Horticulturae,
Fang S, Lang T, Cai M & Han T (2022) Light keys open locks of plant
272:109508.
photoresponses: A review of phosphors for plant cultivation LEDs.
Journal of Alloys and Compounds, 902:163825. R Core Development Team (2021) R: A Language and Environment for
Statistical Computing. R Foundation for Statistical Computing. Avail-
Ferreira EB, Cavalcanti PP & Nogueira DA (2021) ExpDes: Experimen-
able at: <https://www.r-project.org/>. Accessed on: March 15th, 2022.
tal Designs Package. R package version 1.2.2. Available at: <https://
CRAN.R-project.org/package=ExpDes>. Accessed on: March 15th, Sæbø A, Krekling T & Appelgren M (1995) Light quality affects photo-
2022. synthesis and leaf anatomy of birch plantlets in vitro. Plant Cell, Tissue
and Organ Culture, 41:177-185.
Fukuda N, Ajima C, Yukawa T & Olsen JE (2016) Antagonistic action of
blue and red light on shoot elongation in petunia depends on gibberel- Santos RP, Cruz ACFD, Iarema L, Kuki KN & Otoni WC (2008) Proto-
lin, but the effects on flowering are not generally linked to gibberellin. colo para extração de pigmentos foliares em porta-enxertos de videira
Environmental and Experimental Botany, 121:102-111. micropropagados. Revista Ceres, 55:356-364.
Frank HA & Cogdell RJ (1996) Carotenoids in photosynthesis. Photo- Shin KS, Murthy HN, Heo JW, Hahn EJ & Paek KY (2008) The effect
chemistry and Photobiology, 63:257-264. of light quality on the growth and development of in vitro cultured
Doritaenopsis plants. Acta Physiologiae Plantarum, 30:339-343.
Guo X, Hao X, Khosla S, Kumar KGS, Cao R & Bennett N (2016) Effect
of LED interlighting combined with overhead HPS light on fruit yield Wellburn AR (1994) The spectral determination of chlorophylls a and
and quality of year-round sweet pepper in commercial greenhouse. b, as well as total carotenoids, using various solvents with spectro-
Acta Horticulturae, 1134:71-78. photometers of different resolution. Journal of Plant Physiology,
144:307-313.
Gupta SD & Jatothu B (2013) Fundamentals and applications of
lightemitting diodes (LEDs) in in vitro plant growth and morphogene- Wickham H (2016) ggplot2: Elegant Graphics for Data Analysis.
sis. Plant Biotechnology, 7:211-220. Springer, New York. 260p.

Rev. Ceres, Viçosa, v. 70, n. 3, p. 40-53, may/jun, 2023

 Light sources indoor benefit the growth and development of pepper cultivars 53

Zhang X, Bian Z, Yuan X, Chen X & Lu C (2020) A review on the

effects of light-emitting diode (LED) light on the nutrients of sprouts
and microgreens. Trends in Food Science & Technology, 99:203-216.
Zhang YT, Zhang YQ, Yang QC & Li T (2019) Overhead supplemental
far-red light stimulates tomato growth under intra-canopy lighting
with LEDs. Journal of Integrative Agriculture, 18:62-69.
Yang X, Xu H, Shao L, Li T, Wang Y & Wang R (2018) Response of
photosynthetic capacity of tomato leaves to different LED light wave-
length. Environmental and Experimental Botany, 150:161-171.

Rev. Ceres, Viçosa, v. 70, n. 3, p. 40-53, may/jun, 2023