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Photomorphogenesis of wheat sprouts with LED irradiation of different

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Article  in  Lighting Research and Technology · December 2019

DOI: 10.1177/1477153519894716

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Lighting Res. Technol. 2020; 52: 583–594

Photomorphogenesis of wheat sprouts

with LED irradiation of different
intensities
T Han PhDa,b, T Astafurova DScc, S Turanov PhDa , A Burenina MScc, A Butenkova MScc,
E Surnina MScc and D Valiev PhDa,b
a
National Research Tomsk Polytechnic University, Tomsk, Russia
b
Chongqing University of Arts and Sciences, Chongqing, China
c
National Research Tomsk State University, Tomsk, Russia

Received 26 September 2019; Revised 15 November 2019; Accepted 23 November 2019

Definition of the growth and development characteristics of plants in varied light

conditions is a key factor for the creation of highly efficient light facilities for plant
cultivation. Experimental research was conducted using an LED irradiation facility
with photosynthetic photon flux densities ranging from 0 to 261 lmol m2 s1 and
a continuous spectrum with maxima at 445 and 600 nm. Under the maximum
photosynthetic photon flux density (261 lmol m2 s1) wheat germs demonstrated
diminishing leaf surface with high values of specific leaf area, enhanced
pubescence of ground tissues, increases in the number of stomata on the upper
epidermis and palisade, and an increase in the thickness of the leaves as well as an
increase in carotenoids but a decrease in the chlorophyll aþb/carotenoids relation.
It was revealed that the optimum level of photosynthetic photon flux density for
the referred spectrum was in the range from 82 to 100 mmol m2 s1, which may
enable a reduction of irradiance under specific conditions during early develop-
ment with no harm to the plants while minimizing energy consumption during
cultivation.

1. Introduction parameters, may be the most promising for

The modernization of irradiation facilities
and the creation of new highly efficient light
sources are decisive conditions for the devel-
opment of photoculture.1–4 They will ensure
the necessary intensity and optimal ratio of
photosynthetically active radiation (PAR) for
individual spectral regions. In addition, they
will correspond to the operational, technical
and economic conditions of production and
will not require a radical reconstruction of
the premises. As the lighting devices of
the new generation, LEDs, for a number of
Address for correspondence: S Turanov, National Research
Tomsk Polytechnic University, Lenin Avenue 30, 634050
Tomsk, Russia.
E-mail: tyrsb@yandex.ru
use in green planting.5–10
In recent years, the interest of researchers
in this scientific field has increased signifi-
cantly. Numerous reports provide a large
array of data on a variety of LED sources,
different spectral characteristics, luminous
flux intensities, lighting modes, taking into
account the species, varietal and age charac-
teristics of plants, as well as their energy
efficiency, durability, high manufacturability
and profitability. The effect of photon flux on
the structural and functional parameters of
plants depending on their developmental
stage is discussed. It is noted that the different
lighting conditions can regulate the chemical
composition, nutritional value, production
of regeneration material in vitro culture,

ß The Chartered Institution of Building Services Engineers 2019 10.1177/1477153519894716

584 T Han et al.
transplant survival, timing and duration of
flowering, crop productivity, as well as post-
harvest storage of fruits and removal of seeds
from dormancy.1,5 Despite the large amount
of data on the use of LED lighting in crop
production, many questions remain unan-
swered. This makes it difficult to develop a
unified concept and create a common model
for application. Thus, research results that are
optimal for growing plants in closed life-
support systems with full artificial light (e.g.,
in phytotrons or in orbital stations) differ
from the lighting conditions of the same
plants in natural light, which, in turn, are
subject to both climatic and seasonal
changes.9 In the regulation of lighting condi-
tions, it is necessary to take into account the
nature of the economic conditions necessary
for photoculture to significantly expand. The
idea of vertical farming is actively developing,
where, along with traditional green crops,
berry, essential oil, medicinal and other non-
traditional crops are cultivated. The growing
of microgreens, the range of which is con-
stantly expanding, is becoming increasingly
popular for functional nutrition. Microgreens
are young edible sprouts from seeds of
various types of vegetable, grain, aromatic
and wild edible plants.11,12 Depending on the
species, they can be harvested 7–21 days after
germination, when the cotyledon leaves are
fully formed and the first leaves appear.13 The
response of these specialized early crops to
lighting parameters differs from the response
of the same species grown for other economic
purposes.10 Wheat is one of the most common
microgreen species. The young green sprouts
of wheat are used as primary produce for the
manufacture of healthy foods, usually in the
form of juice (wheatgrass) and powder from
dried sprouts. Wheatgrass juice contains most
of the vitamins, macronutrients, enzymes and
amino acids. It is well known that wheat
sprouts contribute to the normalization of
metabolism, increase immunity and eliminate
toxins from the body.14
Lighting Res. Technol. 2020; 52: 583–594
It is difficult to define general irradiation
patterns for various types of microgreens,
since plants with different life strategies may
have special requirements for lighting even at
early stages of development. It is worth
noting that most researchers study the effects
of the spectral composition of radiation on
the growth, quality, and presentation of
microgreens.15–18 It is known that the spectral
composition of light most effectively performs
a regulatory function in the range of low
irradiance levels.19 Therefore, a comparative
study of the effect of light of different
intensities on the anatomical, morphological
and physiological parameters of various types
of microgreen is of considerable interest. The
aim of the present work is to determine the
optimal level of LED irradiation for wheat
seedlings using various parameters of
morphogenesis.
2. Materials and methods
2.1 Plant material and growth conditions
The spring soft wheat (Triticum aestivum
L.) cultivar ‘‘Iren’’ was chosen as a model
plant. The plants were purchased from
Siberian Research Institute of Agriculture
and Peat, Tomsk, Russia. Seed germination
was not less than 95%. Experimental samples
of spring soft wheat were previously sprouted
for two days before the seedling emergence.
The samples were evenly separated into vege-
tation pots with peat (15 g per pot). The size
of the pot was 685.5 cm. The pots were
placed in cells with a microclimate and
irradiation control – phytotrons (Figure 1(a)
and (b)). A control was performed with a
darkened cell of the climate chamber.
Throughout the experiment (10 days), the
temperature of 23  2 8C and humidity 70%
were kept constant in all cells. The plants
were watered manually every day with similar
amount of distilled water for each pot.
 Photomorphogenesis of wheat sprouts 585

(a) (b)

Power supply Electrical current (A),

Driver voltage (V)

LEDs
Lights control

RD 485 Communication
unit

Power supply
Temperature Humidity

CO2 Watering
Climate control

Temperature Humidity CO2 PPFD

Sensors

(c) (d)
Irradiance, mW/m2

40

20

0
400 500 600 700 800
Wavelengh, nm
0 49 128 192 260
µmol·m–2·S–1

Figure 1. (a) Block diagram of the control system; (b) picture of the ‘‘Phytotron’’ greenhouse; (c) specialized irradiation
unit; (d) spectrum of the LED irradiation system

Lighting Res. Technol. 2020; 52: 583–594

586 T Han et al.
Table 1 The parameters of the phytotron irradiation unit
Sample Sample
number 1 number 2
Power consumption of the IU, (W) 0 5
Irradiance, (W/m2) – 11.3
Photosynthetic photon flux – 53.4
density (lmol  m2  s1)
2.2 Intelligent lighting system
The light flux was provided by a specialized
irradiation unit (IU) (Figure 1(b) and (c)),
which allows for the manual and automatic
control of the spectral composition and
irradiation intensity (through three independ-
ent channels). A control system for the
phytotron was developed for the technical
implementation of automatic lighting.
The IU is a plate with circuit boards
located on one side and cooling radiators on
the other. Each cell of the phytotron contains
20 circuit boards, each of which has nine
LEDs of three colours (blue, (Cree
XTEARY), red (Cree XPEPHR) and white
(Cree XTEAWT). The adjustment of irradi-
ation modes was carried out by changing the
current through the LEDs from 0 to 700 mA.
The photon flux of red LEDs can vary from
0 to 264 mmol/s, for white LEDs from 0
to 201 mmol/s, for blue LEDs from 0 to
263 mmol/s. In this way, it is possible
to form practically any spectral composition
in the PAR region and vary the photosyn-
thetic photon flux density (PPFD) in both
automatic and manual control mode. The
design of the phytotron ensures uniform
irradiance on the plant’s surface (Figure 1(c)).
The main elements of the IU are the RS-
485 level converter, an 8-bit AVR microcon-
troller (ATtiny2313-20S), a pulse current
driver LDD-700H, and an electrolytic cap-
acitor battery to form a direct current output.
Software control of load current is carried out
via RS-485 interface (Figure 1(a)).
Lighting Res. Technol. 2020; 52: 583–594
Sample Sample Sample Sample
number 3 number 4 number 5 number 6
10 15 25 50
17.3 20.6 37.7 54.9
82.4 97.4 178.6 261.0
The operating modes of irradiation were
selected in such a way that the total flux of
light in the PAR region was the same in
all cells of the phytotron (continuous spectra
with maxima at 445 and 600 nm) (Figure 1(d)).
This choice of the spectrum is due to the
presence of all components of light in the PAR
region. The total flux of light in the PAR
region for each cell was selected differently
(Table 1). The photoperiod was fixed at 16
hours. The adjustment of irradiation modes
was carried out by changing the current to the
LEDs. This was the most convenient and
easily controlled method for controlling the
intensity of the LED’s radiation (Figure 2).
2.3 Morphometric parameter estimation
Morphometric parameters such as the
length of the root system sprouts, the mass
of the root and aboveground parts of plants
were evaluated to assess the effect of the
irradiation parameters on the vegetation pro-
cesses of plants in all the tested plant samples.
For leaf area the leaves were scanned, using
the program AxioVision (Carl Zeiss). The
number of pixels measured sheet area and the
measurements were translated into square
centimetres. Specific leaf weight (SLW) was
defined as the total dry weight of leaves per
unit of leaf area (with units of mg/cm2).
The anatomical structure of the leaves was
studied in accordance with generally accepted
methods,20,21 the stomatal index (SI) was
calculated by SI (%) ¼ (S/SþE)100, where
S and E are the number of stomata and
 Photomorphogenesis of wheat sprouts 587

epidermal cells, respectively, per mm2.22 Fresh sample, at least 25 sections were analysed.
leaves were prepared by cutting on an MZ – 2 Photographs of microscopic specimens of
freezing microtomes (slice thickness leaves and microscopic measurements were
90–160 mm) in the middle part of the leaf. estimated on a Carl Zeiss Axio Lab A1 optical
The slices were made fivefold on leaves microscope with an AxioCam ERc 5s digital
collected from five plant samples. For each camera using Axio Vision 4.8 Software.
The amount of the pigments was deter-
mined in ethanol (96 %) extracts.
2.4 Absorbances at 665, 645 and 440.5 nm were
Irradiation intensity, [a.u.]

Blue
2.0 read (spectrophotometer UV-1601PC,
Red
1.6 White
‘‘Shimadzu Corp.’’, Japan). The concentra-
tion of chlorophylls was then calculated as
1.2 described by Wintermans and de Mots,23 and
0.8 that of carotenoids according to Wettstein.24
0.4
0.0
0 150 300 150 600 3. Results
Current, mA
3.1 Morphometric parameters of wheat plants
Figure 2. The dependence of the irradiation intensity on
the current through the LEDs These studies conducted with wheat
sprouts grown under different intensities

0 53.4 82.4 97.4 178.6 261

mmol*s–1*m–2 mmol*s–1*m–2 mmol*s–1*m–2 mmol*s–1*m–2 mmol*s–1*m–2 mmol*s–1*m–2

Figure 3. The effect of light treatment with different photosynthetic photon flux densities on wheat sprouts

Lighting Res. Technol. 2020; 52: 583–594

588 T Han et al.

(a) 25 (b) 35
a
30 b b b
20

Shoot length, cm
a
Root length, cm

a a a a 25 c c,d
15 20
b
15
10
10
5
5
0 0

(c) 12 (d) 25
a a
a

Dry weight of sprout, mg

a a a a
Dry weight of root, mg

10 a 20
b
8 b
b 15 b
6
10
4

2 5

0 0

(e) 8 (f) 2.5

a
7 a a
a a 2 b
6 b
SLW, mg cm–2
Leaf area, cm2

c c c
c
5 1.5
4 c
3 1
2
0.5
1
0 0
0 53 82 97 179 261 0 53 82 97 179 261
PPFD, µmol m–2 S–1 PPFD, µmol m–2 s–1

Figure 4. The effect of photosynthetic photon flux density (PPFD) on (a) root length, (b) shoot length, (c) dry weight of
root, (d) dry weight of sprout, (e) leaf area and (f) specific leaf weight (SLW) of wheat sprouts. Different letters indicate
differences with p  0.05

showed effects on a number of morphological 261 lmol  m2  s1, the difference from the
parameters. The length of the seedlings’ roots control test sample was 30% (Figures 3 and
grown without irradiation (0 lmol  m2  s1 4(b)).
– control test sample) were 31–36% shorter The PPFD also had an impact on the
than the plant samples grown under higher accumulation of wheat sprout biomass which
levels of irradiation (Figure 4(a)). At the same increased relative to the control in experimental
time, the shoot length decreased with increas- samples at PPFDs of 82–261 lmol  m2  s1
ing PPFD, and the wheat plants were (Figure 4(c) and (d)). Figure 4(e) shows how
more compact. With a maximum PPFD of the PPFD influenced leaf area. The area
Lighting Res. Technol. 2020; 52: 583–594
 Photomorphogenesis of wheat sprouts 589
Table 2 Anatomical characteristics of wheat leaves when grown in lighting conditions of different photosynthetic
photon flux density (PPDF)

Characteristic PPDF (lmol m2s1)

0 82 261
2
Upper epidermis The number of cells per 1 mm 193.78  10.81c 227.20  5.62b 302.08  5.49a
Stomatal density per 1 mm2 41.78  4.12b 49.28  3.05b 59.52  3.00a
Stomatal index, % 17.36  1.24a 17.61  0.80a 16.34  0.67a
Indument density per 1 mm2 36.44  4.44a 17.28  2.43b 28.16  3.10a
Lower epidermis The number of cells per 1 mm2 254.08  10.79a 225.92  3.50b 226.56  4.40b
Stomatal density per 1 mm2 34.56  3.77b 49.28  4.23a 34.56  2.72b
Stomatal index, % 11.62  0.78b 17.51  1.31a 13.05  0.92b
Indument density per 1 mm2 47.36  4.38a 27.52  3.00b 40.32  3.47a
Leaf slice Lamina thickness in vascular 213.53  4.04b 228.22  1.50a 212.85  9.29a, b
bundles (lm)
Lamina thickness (leaf blade) (lm) 130.19  3.64b 152.54  1.83a 124.68  5.08b
Upper epidermis thickness (lm) 17.71  0.45b 18.32  0.28b 24.94  0.72a
Lower epidermis thickness (lm) 18.84  0.39a 20.62  0.81a 21.26  1.15a
Mesophyll thickness (lm) 61.96  4.02c 121.70  2.47a 87.46  2.38b
Palisade mesophyll thickness (lm) 39.65  1.49c 63.98  1.10a 52.22  2.09b
Spongy mesophyll thickness (lm) 45.42  1.49c 69.77  2.14a 55.00  1.43b
Palisade/spongy ratio 0.90  0.05a 0.94  0.04a 1.03  0.09a
Height of upper mesophyll layer 26.74  0.70b 36.62  1.07a 36.98  0.77a
cells (lm)
Width of the upper mesophyll 22.87  0.67a 18.70  0.73b 21.60  0.45a
layer cells (lm)
Chloroplast diameter (lm) – 5.94  0.17a 4.71  0.12b
Vascular bundle cross-sectional 2774.26  183.50b 3886.77  416.74a 3464.03  250.49a
area (lm2)

Different letters indicate differences with p  0.05.

increases by 63% and reached a maximum for
the plant sample at a PPFD of
82 lmol  m2  s1 – 77% more than the con-
trol. With a further increase of PPFD, the area
of formed leaves gradually decreased to a
PPFD of 261 lmol  m2  s1.
An important parameter of the leaf,
characterizing the vital activity of plants, is
the SLW. The SLW is significantly lower than
the control condition at a PPFD of 261 lmol
m2  s1. It is also, in general, significantly
different from the other experimental variants
(Figure 4(f)). A high value of SLW with the
highest intensity of radiation reflects the
accumulation of a larger amount of dry
mass per unit leaf area.
3.2 Anatomical characteristics
A number of key indicators were deter-
mined when studying the anatomical
structure of the leaves of wheat grown under
the lighting conditions of varying intensity.
A significant part of these indicators
had maximum values in wheat leaves
grown at a PPFD of 82 lmol  m2  s1: The
thickness of the leaves, mesophyll and its
components, as well as the diameter of
chloroplasts, the cross-sectional area of the
vascular bundle, the number of stomata and
the stomatal index of the lower epidermis
(Table 2).
Wheat leaves at a PPFD of 261 lmol
m2  s1 were characterized by the greatest
number of cells and stomata of the upper
epidermis, as well as the greatest thickness of
the upper and lower epidermis (Table 2,
Figure 5) when the lamella thinned. This
reflects their adaptation to the conditions of
intense lighting. It should be noted that the
stomatal indices of the upper epidermis do
Lighting Res. Technol. 2020; 52: 583–594
590 T Han et al.
(a) (b)
0 µmol·m–2·s–1
500 µm
Figure 5. Cross-sections of wheat when grown under different photosynthetic photon flux density (PPDF) conditions.
(a) 0 mmol m2 s1; (b) 82 mmol m2 s1; (c) 261 mmol m2 s1
not have significant differences at PPFDs of
0, 82 and 261 lmol m2  s1.
3.3 The content of photosynthetic pigments
The main acceptor of light is the pigment
system of plants, which reacts to the amount
and spectral composition of the incident light,
that later has a great influence on photosyn-
thetic productivity.25 It was found that in the
control condition (PPDF of 0 lmol  m2  s1,
without light), the lowest photosynthetic pig-
ment concentration in the leaves was
observed, and the maximum amount of
chlorophyll a and b per leaf area was observed
at a PPFD of 82 lmol  m2  s1. It should be
noted that the content of chlorophyll a and
carotenoids do not significantly differ
between PPFDs of 82–179 lmol  m2  s1
(Table 3).
The amount of carotenoids increases at the
maximum PPFD (261 lmol  m2  s1) by
8–14% compared with the other light vari-
ants. There was a tendency to increase the
chlorophyll a/b ratio and a significant
decrease in the chlorophyll a þ b/carotenoids
relation when exposed to high PPFD.
4. Discussion
The most important characteristics of the
LED illumination mode for plants are the
Lighting Res. Technol. 2020; 52: 583–594
(c)
82 µmol·m–2·s–1 261 µmol·m–2·s–1
500 µm 500 µm
photosynthetic photon flux density (PPFD) in
the spectral range of photosynthetically active
range, the photoperiod, the spectral compos-
ition of radiation and the mode of illumin-
ation (constant or variable). The range of
PPFD values that stimulate plant growth
is usually from 200–250 lmol  m2  s1 to
500–800 lmol  m2  s1.9 In greenhouses,
leafy vegetables are usually grown with PPFD
levels of 200–300 lmol  m2  s1,23 but there
are data indicating that lettuce is grown
at PPFD levels of 300–400 lmol  m2  s1.24
For Chinese cabbage, a PPFD of
672 lmol  m2  s1 is considered saturating.25
When grown at a PPDF of more than
600 lmol  m2  s1, in some cases, the photo-
synthetic apparatus of the upper leaves and the
total yield biomass decrease as compared to
plants grown at lower levels of PPFD.26
In recent years, there has been considerable
interest in the production of microgreens,
which constitutes a growing market segment
in the crop production sector. A number of
studies have shown that different levels of
PPDF have been used for growing micro-
greens of different crops: for cabbage –
50 lmol  m2  s1, basil – 170 lmol  m2  s1,
lettuce – 135 lmol  m2  s1, white mustard,
spinach, parsley and green onions –
300 lmol  m2  s1.10,30 Thus, it is difficult to
establish a single optimal level of PPFD – both
in terms of the needs of plants, and the
 Photomorphogenesis of wheat sprouts 591
Table 3 Effect of photosynthetic photon flux density on the photosynthetic pigments content of the first wheat leaf,
(mgdm2)

Photosynthetic photon Chl a Chl b Chl a / Chl b Carotenoids Chl aþb/

flux density (lmol m2s1) carotenoids

0 (reference material) 0.03  0.003c 0.01  0.001c 5.3c 0.11  0.01d 0.3c
53 1.38  0.05b 0.44  0.02b 3.1b 0.34  0.01c 5.3b
82 1.52  0.07a 0.51  0.03a 3.0b 0.39  0.01b 5.3b
97 1.46  0.03a 0.46  0.01b 3.2b 0.37  0.01b 5.3b
179 1.44  0.03a 0.46  0.01b 3.1b 0.37  0.01b 5.2b
261 1.44  0.04a 0.43  0.01b 3.4a 0.42  0.01a 4.5a

Different letters indicate differences with p  0.05.

Chl a: chlorophyll a, Chl b: chlorophyll b, Chl a/Chl b: chlorophyll a/chlorophyll b, Chl a þ b/carotenoids: (chlorophyll
a þ chlorophyll b)/carotenoids.

profitability of their production. photosynthetically active tissues in relation

Recommended levels of PPFD depend on the
type of plant, planting density, growth condi-
tions, and on the specific cultivation target.
This may be the maximum productivity of the
shoot (biomass) for leafy vegetables, maximum
seed productivity for grain crops, rapid or
simultaneous flowering of flower crops, etc.
For most agricultural plants, even a small
increase or decrease in light intensity leads to
significant changes in morphology and anat-
omy,31 as well as the rate of photosynthesis,
transpiration and stomatal conductance of
leaves.32,33 According to our investigations,
the highest biomass indices were observed
in experimental samples with PPFD of
82–261 lmol  m2  s1. We did not detect
any differences in the given indicator in this
irradiation range. The leaf area practically
do not change over a PPFD range of
53–179 lmol  m2  s1 and decreased at a
PPFD of 261 lmol  m2  s1. The SLW
values have an inverse pattern with a decrease
in the average range and an increase at
maximum PPFD. It is believed that in
plants with low SLW i.e., with a thinner
leaf, the photosynthesis is carried out more
efficiently due to lower costs for the leaf
formation and loss of light energy. At the
same time, a decrease in SLW value may
reflect an increase in the share of
to the volume of cell walls in the dry mass of
leaves,34 which was noted in the samples with
a PPFD of 82 lmol  m2  s1. The increase
of SLW at the maximum PPDF of
261 lmol  m2  s1 was due to an increase in
the upper and lower epidermis thickness with
a decrease in the leaf plate thickness.
It is well known that with a higher PPFD,
both the leaf thickness and the palisade
and spongy mesophyll increase.35–37 In
our experiment, an increase in these charac-
teristics was observed at a PPFD of
82 lmol  m2  s1, and at higher levels
of exposure (261 lmol  m2  s1), a decrease
in these parameters was observed, which is
consistent with the results of studies con-
ducted for other plant species.35–38
Analysis of the content of photosynthetic
pigments in the leaves of the experimental
samples revealed their minimum concentra-
tion in a low PPFD (53 lmol  m2  s1). As
the PPFD increased (up to
261 lmol  m2  s1), an increase in the con-
centration of chlorophylls and carotenoids
was observed, without significant differences
between the experimental samples. With a
high PPFD (261 lmol  m2  s1), a decrease
in the chlorophyll content was observed,
which is apparently associated with an
increased degeneration of these pigments.39–41
Lighting Res. Technol. 2020; 52: 583–594
592 T Han et al.
5. Conclusion
The experimental data show that at a PPFD
of 82 lmol  m2  s1 wheat sprouts show the
maximum leaf parameters such as the leaf and
mesophyll thickness, the size of chloroplasts,
photosynthetic pigment content, and leaf
area. In plants grown in the PPFD range
from 82 to 179 lmol  m2  s1 there are no
differences in such indicators as weight and
length of root, aerial parts weight, SLW, and
leaf area. These obtained results allow us to
conclude that the yield and market quality of
wheat sprouts are the best for samples grown
at a PPFD of 82 lmol  m2  s1.
Thus, the optimal PPFD with a continuous
spectrum of LED radiation with maxima at
445 and 600 nm is in the range from 82 to
100 lmol  m2  s1. This will make it possible
to reduce the irradiation in the early stages of
development without harm to the plants
which will reduce the energy consumption
required for their cultivation.
Acknowledgement
We are very grateful to Alexey Yakovlev for
their essential support that made this research
possible.
Declaration of conflicting interests
The authors declared no potential conflicts of
interest with respect to the research, author-
ship, and/or publication of this article.
Funding
The authors disclosed receipt of the following
financial support for the research, authorship,
and/or publication of this article: This research
was supported by Tomsk Polytechnic
University CE Program and Tomsk State
Lighting Res. Technol. 2020; 52: 583–594
University competitiveness improvement pro-
gram grant (no. 8.1.29.2018).
ORCID iD
S Turanov https://orcid.org/0000-0002-
9518-0598
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