Scientia Horticulturae 250 (2019) 81–88

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Scientia Horticulturae
journal homepage: www.elsevier.com/locate/scihorti

Effects of daytime intra-canopy LED illumination on photosynthesis and T

productivity of bell pepper grown in protected cultivation
Naveen C. Joshia,1, Kira Ratnera,1, Orly Eidelmana, Dominika Bednarczyka, Naftali Zura,
⁎
Yair Manya, Yosepha Shahaka, Elinor Aviv-Sharonb, Meir Achiamc, Ziva Giladc, Dana Charuvia,
a
Institute of Plant Sciences, Agricultural Research Organization (ARO) - Volcani Center, Rishon LeZion, 7505101, Israel
b
The Robert H. Smith Institute of Plant Sciences and Genetics in Agriculture, Hebrew University of Jerusalem, Rehovot, 7610001, Israel
c
Jordan Valley Research and Development Authority, Mobile Post, 91906, Israel

A R T I C LE I N FO A B S T R A C T

Keywords: In the past decade, light-emitting diodes (LEDs) have been replacing most other types of light sources. One
Light-emitting diodes (LEDs) emerging use for LEDs in horticulture is ‘intra-canopy illumination’ or ‘LED-interlighting’, feasible owing to their
Intra-canopy illumination low heat output and small physical size. The interlighting technique, typically implemented in environmentally-
Interlighting controlled greenhouses also supplemented with overhead lighting, is mostly relevant for high-wire vegetable
Bell pepper
cultivation, in which self-shading results in light limitations for a large fraction of the canopy. Interlighting has
Photosynthesis
been shown to increase the yield and/or improve fruit quality in all-year round greenhouse crops such as tomato,
Fruit-yield
cucumber and sweet pepper. In this work, we utilized daytime supplemental intra-canopy LED illumination for
sweet pepper grown in high-density ‘Spanish’ trellis systems within passive high tunnels in the Jordan Valley,
Israel (latitude ˜32 °N). While canopy top at these conditions is not light-limited, extensive deep shading of the
inner canopy is a disadvantage. In two experiments carried out in two separate seasons, the supplemental
lighting, which enhanced the photosynthetic rates of the inner canopy foliage by 3.5- to 5.7-fold, resulted in
significant increase (˜30%) of the fruit yield during the spring season. The added yield was attained by higher
fruit numbers, with no notable effects on fruit size or weight. Our results raise the prospect that LED-interlighting
may be a useful practical tool for maximizing fruit production, even in geographical regions of ample sunlight.

1. Introduction
In recent years, with the advancements in the light-emitting diode
(LED) technology, the use of LED lighting in horticulture has been
steeply rising (Bantis et al., 2018; Mitchell, 2015; Mitchell et al., 2015;
Morrow, 2008; Nelson and Bugbee, 2014). As LEDs are essentially solid-
state semiconductor devices, they are able to exhibit very cool photon-
emitting surfaces that generate low heat outputs (Singh et al., 2015).
This characteristic, coupled with the fact that LED chips are quite small
in size, allows using them as supplemental lighting within plants’ ca-
nopies without overheating or burning the plant tissues (Gómez et al.,
2013). Application of supplemental illumination within the canopy is
coined ‘(LED)-interlighting’ or ‘intra-canopy illumination’ (Mitchell
et al., 2015), used here interchangeably. The effects of LEDs, as top- or
inter-lighting, on greenhouse crops can be numerous. Some of the
beneficial ones include added crop-yield, higher quality yield, manip-
ulation of harvest dates, less yield fluctuations during the growth cycle,
as well as enhanced nutritional value (Carvalho et al., 2016; Chen et al.,
2016; D’Souza et al., 2015; Folta and Childers, 2008; Li and Kubota,
2009; Massa et al., 2008; Olle and Virsile, 2013; Ouzounis et al., 2015;
Singh et al., 2015).
LED-interlighting has been mostly studied and utilized in northern
countries (e.g., the Netherlands, Finland, Canada and northern areas of
the USA) in combination with overhead illumination, required due to
the low irradiance and short day length, especially in winter (Dorais,
2003; Hemming, 2011). In these regions, illumination is typically ap-
plied in the context of actively-controlled greenhouses (e.g., Gómez and
Mitchell, 2014; Hao et al., 2012). Intra-canopy LED illumination is most
commonly implemented for year-round high-wire greenhouse vege-
table cultivation, i.e., for cucumber, tomato and pepper (Gómez and
Mitchell, 2014; Jokinen et al., 2012; Trouwborst et al., 2010) [for
earlier interlight studies performed with high-pressure sodium (HPS)
lamps see Gunnlaugsson and Adalsteinsson, 2006; Hovi et al., 2004].
Insufficient light within the canopy of such intensively-grown crops

⁎
Corresponding author.
E-mail address: charuvi@volcani.agri.gov.il (D. Charuvi).
1
These authors contributed equally to the work.

https://doi.org/10.1016/j.scienta.2019.02.039
Received 21 September 2018; Received in revised form 7 February 2019; Accepted 13 February 2019
Available online 19 February 2019
0304-4238/ © 2019 Elsevier B.V. All rights reserved.
N.C. Joshi, et al.
limits photosynthesis, flowering and/or fruit set, and thus may limit the
productivity.
Bell pepper is known to exhibit high rates of organ abortion, whe-
ther of flower buds, flowers or young fruit, with different environ-
mental factors affecting this phenomenon (Wubs et al., 2009). The ef-
fects of light on abortion have been studied mostly with relation to light
intensity and photoperiod. Shading experiments have shown that
flower and/or bud abortion rates decrease at higher light intensities, in
an intensity-dependent manner (Aloni et al., 1996; Shifriss et al., 1994).
Extending the photoperiod likewise has the potential to increase the
yield by increasing the number of fruits (Demers et al., 1998). Studies
characterizing the sink and source strengths have indicated that both
play cardinal roles in the determination of flower and fruit abortion
(Aloni et al., 1997; Marcelis et al., 2004; Turner and Wien, 1994).
However, the distinct sensitivities of different pepper cultivars to shade-
induced abscission could not be attributed to photosynthetic rates per se
(Aloni et al., 1996; Turner and Wien, 1994). This is albeit the fact that
leaf photosynthesis is a major determinant of the sink-source relation-
ship and assimilate partitioning. Furthermore, González-Real et al.
showed that leaf photosynthetic characteristics in pepper are affected
more predominantly by the adjacent fruit sink demands rather than just
light acclimation (González-Real et al., 2009). Nonetheless, it may be
expected that at extremely low light conditions, as may be the case
within inner shaded canopies, the very low assimilation rates of the
leaves render them a weak source, leading to the abscission of nearby
organs.
Characterization of the light response curve of leaf photosynthesis
along canopy height in sweet pepper revealed that the contribution of
the bottom half of the canopy to net photosynthesis, estimated using
light photosynthetic parameters and a model of light distribution within
the canopy, was negative (−0.5%) on a yearly basis (Dueck et al.,
2006). Correspondingly, it was shown that pepper plants benefit from a
more uniform distribution of light within the canopy: an increase of
14% in the photosynthetic photon flux (using HPS as the light source) at
different canopy heights resulted in an increase of 23% in the total fruit
yield (Hovi-Pekkanen et al., 2006). The use of photoselective nets that
possess light-scattering properties likewise resulted in significant in-
creases of yield in pepper owing to better penetration of light into the
inner canopy (Shahak, 2008).
In the Jordan Valley, Israel, a region with ample sunlight (see DLI,
Fig. 1B), bell pepper is commonly grown in double-row beds applying
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Scientia Horticulturae 250 (2019) 81–88
the ‘Spanish’ trellising system, within passive walk-in tunnels (Fig. 1A).
In this growth regime, canopy height develops quite rapidly, resulting
in extensive shaded regions of the canopy at the center of the beds
during most of the season. The outer parts of the canopy are, however,
mostly not light-limited. In this work, we set to determine whether
supplemental intra-canopy illumination can be utilized for increasing
the fruit yield of bell pepper at these (variable) conditions, namely
within passive tunnels. This is as opposed to most interlighting studies,
which have been conducted within actively-controlled greenhouses.
The supplemental LED illumination was applied during the daytime
hours starting ˜2.5 months after planting and until the end of the
season. In two experiments conducted during two growth seasons, we
found that the intra-canopy illumination, which significantly enhanced
the photosynthetic rates of the inner canopy foliage, increased the fruit
yield by ˜30% in the spring months. The increase in yield was realized
by a higher number of fruits, yet without affecting fruit size or weight.
2. Materials and methods
2.1. Plants and growth conditions
The study was conducted at the Zvi R&D Experimental Station in the
Jordan Valley (‘Mop-Bika’, 31°59′49.0″N, 35°27′09.3″E) in a high
plastic tunnel (10 m wide x 4 m high x 45 m long) in the local soil, a
well-drained (EC < 2.0 dS/m) clay (30%) – limestone (50%) marl soil.
Management during the experiments followed the routine practices of
the region. Prior to planting, soil sanitation was carried out by solar
fumigation for 4 weeks, in the last week of which metam sodium
(40 mL/m2) was streamed into the soil via the drip irrigation system.
Fertilization was carried out based on soil sampling, using N-P-K
(6:3:9), with a total seasonal nitrogen content of ˜50 g/m2. Experiments
were carried out during two growth seasons: from Aug. 2016 to the end
of Apr. 2017 (2016–2017) and from Aug. 2017 to the end of Apr. 2018
(2017–2018). Red bell pepper seedlings (Capsicum annuum L. cv.
Cannon, Zeraim Gedera/Syngenta) were planted on Aug. 8th (2016) /
Aug. 14th (2017) in double-row beds. The width of the double-row beds
was 0.8 m, and the center-to-center distance between beds was 1.75 m,
with an overall plant density of 2.9/m2. Plant training applied the
‘Spanish’ trellis system, with lateral horizontal wires supporting the
canopy vertically, and without any pruning. In this growth system,
canopy height reaches 2–2.2 m. At the time of planting, the tunnel was
Fig. 1. Bell pepper growth in the Jordan
Valley. (A) Bell pepper grown in double-row
beds applying the ‘Spanish’ trellising system in
a passive walk-in tunnel at the Jordan Valley
Experimental Station. Image was taken on Jan.
30th, 2017. (B) Monthly mean daily light in-
tegral (DLI) values ± SD during the months of
two experiments (Aug. to end of Apr.,
2016–2017 and 2017–2018), calculated from
solar radiation data collected by the Israeli
Meteorological Service at the Gilgal Station.
Dashed lines indicate transmission into the
tunnel, which varies in the different months
according to the tunnel coverings, as detailed in
the text. (C and D) Daily minimal (T min, blue)
and maximal (T max, red) temperatures from
data collected at Gilgal for the time periods in
which the experiments were conducted in
2016–2017 (C) and 2017–2018 (D). (For in-
terpretation of the references to colour in this
figure legend, the reader is referred to the web
version of this article).
N.C. Joshi, et al.
covered by a 50 mesh insect-proof screen, as well as a black shade net
(40%). On Sept. 8th (2016) / Sept. 11th (2017), the shade net was
removed, and on Nov. 8th (2016) / Nov. 2nd (2017) the mesh screen
was replaced by a polyethylene sheet (Kritifil TUV 3410 CE, Plastika
Kritis, S.A., Iraklion, Greece). To prevent fruit heat damage, on Feb.
23rd (2017) / Mar. 1 st (2018), the plastic sheet was removed and the
50 mesh screen as well as the black shade net (40%) were replaced on
top of the tunnel until the end of the season. Fruits were harvested
according to the commercial standard of picking at 60% red color,
which in winter is every 2–3 weeks and in spring every 1–2 weeks.
Fruits were sorted according to size and quality: ‘class 1′ fruit are ex-
port-quality ones (in all size categories), while ‘class 2′ represent mis-
shapen fruits, or ones with chilling injuries or blossom-end rot.
2.2. Supplemental illumination
The supplemental LED illumination was home-made, based on off-
shelf LED strips (9.6 W/m) purchased from LEDART led solutions
(Hagor, Israel). Spectra of LED strips used to construct the fixtures were
warm-white (‘WW’), cool-white (‘CW’), red (‘R’, 660 nm), blue (‘B’,
455 nm) and green (‘G’, 520 nm). In the first experiment (2016–2017),
four spectral compositions (WW, CW, RB, RGB, Fig. 2) were applied,
each along a 5-m section of the bed. Three sections of the same length
served as controls. In the second experiment (2017–2018), cool-white
(CW) LED fixtures were applied along 5-m sections with 3 replicates,
and 6 sections of the same length served as controls. The fixtures were
installed facing upwards between the two adjacent rows of the central
beds (Fig. 3A). Two identical LED fixtures were used for each section, at
two heights within the canopy. The illumination period began ˜2.5
months after planting, on Nov. 1 st (2016) / Oct. 26th (2017), with
fixtures placed 80- and 120 cm above ground level. On Dec. 26th
(2016) / Dec. 12th (2017) the fixtures were raised up to 100- and
140 cm above the ground to follow the height of the productive regions
of the plants. Spectra and photosynthetic photon flux densities (PPFD)
were recorded using a portable spectroradiometer (EPP2000C, Stel-
larNet, Inc., USA) with a cosine−corrected head (Apogee Instruments
Inc., USA) and an LI-250 A quantum sensor (LI−COR, USA), directly
above the light fixtures at a distance of 10 cm (Fig. 2). Illumination was
applied during the natural daylight hours (adjusted accordingly along
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Scientia Horticulturae 250 (2019) 81–88
the season, Supp. Table S1). Leaf temperature was measured using an
infrared thermometer (AZ Instrument Corp., Taiwan) at a distance of
20 cm from the leaves. The air temperature was recorded using EC650
MicroLogs (Fourtec - Fourier Technologies, Israel) hung in proximity
(10–15 cm) to the LED fixtures or at the same heights in control sec-
tions.
2.3. Chlorophyll content and fluorescence
Chlorophyll content was measured non-destructively in attached
leaves using a SPAD-502 chlorophyll meter (Spectrum Technologies,
Inc., USA). Chl-a fluorescence (Fv/Fm), was measured using a portable
pulse-amplitude-modulated (PAM) fluorometer (PAM-2000, Heinz
Walz GmbH, Germany) in attached leaves following a 20-min dark
adaptation with dark clips.
2.4. Gas-exchange measurements
Gas-exchange measurements were performed on attached mature
leaves of the inner canopy. In the LED-illuminated plots, leaves were
sampled 10–25 cm above the lower LED fixtures - at a height of
110–120 cm above the ground, and at the same height in control (non-
illuminated) plots. Measurements were conducted between 9:30 and
11:30 a.m. using a portable LCi photosynthesis system (ADC
BioScientific Ltd., UK) with a clear top chamber at ambient conditions.
Relative air humidity in the tunnel was 62 ± 14%. Leaf temperature
ranged between 27 to 31 °C for measurements conducted on different
dates, with no significant differences between control and LED-illumi-
nated leaves on each specific date. The positioning of the LCi chamber
was according to the orientation of the leaf being probed (in order to
keep the leaf attached), as well as to the light affecting the leaf. In
control plots, the chamber was mostly facing upward (and at an angle
corresponding to leaf orientation) to account for any incoming light
from above. In LED-illuminated plots, the leaf chamber was directed
toward the light originating from the LED fixtures, and, again, main-
taining the original orientation of the leaf. Photon flux density incident
at the leaf level, as recorded by the LCi during the measurements,
averaged at 20 μmol m−2 s−1 or 100 μmol m−2 s−1, respectively for
leaves from control or LED-illuminated plots, at the canopy height
Fig. 2. Spectra of light-emitting diode (LED) fixtures utilized
as intra-canopy illumination. WW, warm white; CW, cool white;
RB, red-blue; RGB, red-green-blue. The photosynthetic photon
flux densities (PPFD) provided by the LEDs used for the first ex-
periment were between 151–162 μmol m−2 s−1 at a distance of
10 cm directly above the fixtures, as denoted on the graphs. CW
fixtures used in the second experiment provided 185 μmol m−2
s−1 at 10 cm.
N.C. Joshi, et al.
Fig. 3. LED illumination setup within the canopy. (A) Two red-green-blue (RGB) LED light segments within the canopy. (B) Side-view of the illumination at the
end of January 2017 (RGB, year 1) and (C) at the end of November 2017 (cool-white, year 2). (For interpretation of the references to colour in this figure legend, the
reader is referred to the web version of this article).
indicated above. For comparison, attached outer-canopy leaves, ex-
posed to natural sunlight in the tunnel were also probed.
2.5. Daily light integral (DLI) calculation
Hourly total solar radiation data (285–2800 nm) for the relevant
months in which the experiments were carried out (Aug. 2016 to Apr.
2017 and Aug. 2017 to Apr. 2018) were obtained from the Gilgal sta-
tion, Israeli Meteorological Service. Daily light integral (DLI) values in
quantum units were calculated by assuming that photosynthetically
active radiation (PAR, 400–700 nm) accounts for 45% (Goldberg and
Klein, 1977; Jacovides et al., 2003) of the total solar radiation, and
using the conversion factor 4.57 μmol m−2 s−1 per W m−2 (Korczynski
et al., 2002; Thimijan and Heins, 1983).
2.6. Statistical analysis
Statistical analysis was performed using one-way analysis of var-
iance (ANOVA), followed by Tukey’s multiple comparisons as a post
hoc test. For comparisons of two groups, the Student’s t-test was used.
The spring yield in the single LED plots of year 1 were compared to the
control using one-sample t-test. All tests were based on a significance
level of 0.05 (unless otherwise noted). Calculations were performed
using the JMP 10 software (SAS Institute, NC, USA). Plots were gen-
erated using MATLAB R2017b (The MathWorks, Inc., Natick,
Massachusetts, USA).
3. Results
3.1. Intra-canopy LED illumination in bell pepper
The study included two experiments carried out during the growth
seasons of 2016–2017 and 2017–2018. These were conducted within a
passive walk-in tunnel (Fig. 1A), and therefore, temperature conditions
during the experiment varied along the season according to the outdoor
ones (Fig. 1C and D). Two and a half months after planting (roughly at
the beginning of Nov.) plant height was already ˜2 m, with the average
light intensity incoming into the canopy from above (between the two
rows of the beds) at a height of ˜1 m being 35 ± 16 μmol m−2 s−1. At
the same height, the intensity encountered in the inner canopy facing
downward was ˜3 μmol m−2 s−1 [these values represent ones obtained
around mid-day on a clear day]. This pronounced self-shading of the
canopy, along with the decrease in night temperature, result in non-
optimal conditions for fruit set. Thus, at this time (2.5 months after
planting) we began applying the supplemental illumination within the
canopy, at the center of the beds.
In the first experiment, four spectral compositions were tested as
intra-canopy illumination: warm-white (WW), cool-white (CW), red-
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Scientia Horticulturae 250 (2019) 81–88
blue (RB) and red-green-blue (RGB) (Fig. 2). These provided
151–162 μmol m−2 s−1 at a distance of 10 cm above the light fixtures.
The second experiment, in the following season, was carried out ap-
plying CW LEDs as intra-canopy illumination, chosen as the spring fruit
yield obtained with this type of LED was the highest in the first season
(see section 3.3). Aiming to achieve improved yield results in the
second experiment, and as no negative effects were noted in the first
experiment, the intensity applied in the second year was raised by
˜15%, providing ˜185 μmol m−2 s−1 at 10 cm (the maximal achievable
with the utilized set-up). In both experiments, two identical 5-m-long
LED fixtures were installed at two heights within the canopy (Fig. 3), in
order to maximize the fraction of inner canopy subjected to the added
light.
Monitoring air temperature in the vicinity of the LED lamps, we
found that compared to the control (non-illuminated) maximal daily
temperature in illuminated areas was higher by 2.4 ± 1.3 and
2.1 ± 0.3 °C along two separate one-month periods recorded (Supp.
Fig. S1). As expected, there were no differences in minimum daily
temperature, as LEDs operated only during the daytime. Differences in
leaf temperature between plants in some of the control and illuminated
plots were also noted; these were mostly on the range of 1–2 °C (Supp.
Table S2) and were not all significant. Thus, the LED illumination did
not result in considerable heating of the inner canopy foliage during the
day.
3.2. Effects of the intra-canopy illumination on photosynthetic parameters
Chlorophyll content and the maximum quantum yield of photo-
system II (Fv/Fm) were surveyed non-destructively in leaves of the
inner canopy with or without the supplementary LED illumination. No
significant differences were found for both parameters for inner canopy
leaves of the control vs. illuminated plants (data not shown).
The effects of the supplemental intra-canopy LED illumination on
photosynthetic rates and stomatal conductance, as measured on at-
tached leaves at the inner part of the canopy are shown in Table 1.
Photosynthesis rates (A, CO2 assimilation) of inner-canopy leaves of
control, non-illuminated, plants were quite low, typically < 1 μmol
CO2 m−2 s−1 (Table 1, ‘CR’). This is in accord with control leaves being
subjected to low light intensities (on average ˜20 μmol m−2 s−1) within
the shaded canopy. Inner-canopy leaves of plants subjected to the
supplemental illumination, at a distance of 10 to 25 cm from the light
fixtures, were subjected to an average intensity of ˜100 μmol m−2 s−1.
These leaves exhibited CO2 assimilation rates which were on average
3.5- to 5.7-fold (on different dates surveyed) higher than control ones.
The effects were similar for leaves illuminated by the various LED fix-
tures utilized in the first and second seasons. Expectedly, the assim-
ilation rates of outer canopy leaves, which are naturally exposed to
higher light intensities, were ˜3-fold higher than those of LED-
N.C. Joshi, et al. Scientia Horticulturae 250 (2019) 81–88

Table 1
Gas exchange parameters of attached inner canopy leaves with or without supplemental intra-canopy LED illumination.
A gs A gs
μmol CO2 m−2 s−1 mol H2O m−2 s−1 μmol CO2 m−2 s−1 mol H2O m−2 s−1

Year 1 Feb. 2017 Mar. 2017

CR 0.71 ± 0.46a 0.015 ± 0.008a 0.90 ± 0.49a 0.028 ± 0.025a
WW 2.52 ± 1.08b 0.084 ± 0.042b 3.05 ± 0.35b 0.053 ± 0.017a
CW 2.87 ± 0.42b 0.103 ± 0.051b 2.77 ± 1.24b 0.053 ± 0.024a
RB 3.10 ± 1.17b 0.052 ± 0.022ab 2.96 ± 1.64b 0.093 ± 0.092a
RGB 2.93 ± 0.72b 0.085 ± 0.042b 3.74 ± 0.53b 0.063 ± 0.019a
Year 2 Jan. 2018 Feb. 2018
CR 1.06 ± 0.77a 0.021 ± 0.011a 0.81 ± 0.41a 0.012 ± 0.009a
CW 3.90 ± 1.14b 0.074 ± 0.029b 4.64 ± 0.81b 0.054 ± 0.017b
sun-exposed 13.45 ± 1.29 0.337 ± 0.092 13.94 ± 3.64 0.435 ± 0.174

Shown are photosynthetic (CO2 assimilation) rates (‘A’) and stomatal conductance to H2O (‘gs’) measured on attached leaves of the inner canopy of control plants
(‘CR’, not illuminated) or of plants illuminated with different LED light fixtureswithin the canopy. WW, warm white; CW, cool white; RB, red-blue; RGB, red-green-
blue. Sun-exposed (outer canopy) leaves were probed at the natural light conditions prevailing in the tunnel (˜760 and ˜640 μmol m−2 s−1 for measurements
conducted in Jan. and Feb. 2018, respectively). Data represent means ± SD. Year 1: n = 4–5 leaves from each plot; Year 2: n = 12 leaves from 2 to 3 plots of CR or
CW each, and n = 4–6 leaves for sun-exposed leaves. Distinct letters (for inner-canopy leaves) indicate statistically significant differences (p < 0.05).

illuminated leaves (Table 1, ‘sun-exposed’). Nonetheless, the differences Table 3

between inner-canopy leaves of LED and CR plants were considerable. Class 1 fruit yield (number m−2) with or without supplemental intra-canopy
Stomatal conductance to H2O (gs) was somewhat higher in inner ca- LED illumination.
nopy leaves of illuminated plants as compared to ones of control plants, Autumn Winter Spring Total
though in year 1 these were not significant for the most part. In the
second experiment, in which more leaf replicates were used, the dif- Year 1
CR 6.4 ± 0.8a 16.0 ± 4.9a 24.7 ± 4.0a 47.1 ± 5.2a
ferences are evident, with LED-illuminated leaves exhibiting ˜3.5- and
LED* 4.7 ± 1.7a 16.2 ± 2.3a 35.9 ± 5.3b 56.8 ± 5.1a
˜4.5-fold higher stomatal conductance than control leaves (Table 1, WW 2.3 18.9 32.5 53.7
Year 2). CW 4.7 14.9 43.0 62.6
RB 6.3 13.8 31.4 51.5
RGB 5.4 17.1 36.8 59.3
3.3. Fruit yield and quality
Year 2
CR 11.8 ± 2.9a 11.5 ± 1.2a 21.1 ± 1.7a 44.3 ± 3.2a
Tables 2 and 3 present the fruit yield obtained in control (CR, non- LED 13.8 ± 0.9a 13.7 ± 1.8a 28.0 ± 3.4b 55.5 ± 6.0b
illuminated) and LED-illuminated plots of the two experiments. In the
first experiment, four different spectral types (WW, CW, RB and RGB) Yield by fruit number represents class 1 fruit and is shown for autumn (Nov.-
were used, each applied to a single 5-m-long section of the bed. As the Dec.), winter (Jan.-Feb.) and spring (Mar.-Apr.). Data represent means ± SD.
Year 1: CR – mean of 3 control, non-illuminated plots; LED* - mean of 4 illu-
light intensity of the different LED types was roughly the same (< 10%
minated plots (WW, CW, RB, RGB). The yield in each single illuminated plot is
difference), and a very similar effect on leaf photosynthesis was ob-
also shown. WW, warm white; CW, cool white; RB, red-blue; RGB, red-green-
served in all types, we compared the average yield in the four illumi- blue. Year 2: CR – mean of 6 non-illuminated plots; LED – mean of 3 plots
nated plots (LED* in Tables 2 and 3) to that of the average CR. This illuminated with CW. Distinct letters (separately for each year and season)
allowed to assess the effect of added light as a basis for the second denote statistically significant differences (p < 0.05).
experiment. For the latter, we chose to apply CW light, as the yield
using this type of LED was the highest in the first experiment (see LED illumination, which started around the beginning of Nov., did not
below). affect the fruit harvested from Nov. to Feb., and is in agreement with
In the first experiment, no significant differences in yield (kg m−2) the fact that the fruit set of this yield occurred prior to the onset of the
between the LED-illuminated and control treatments were found during supplemental illumination. During the spring months, on the other
the autumn and winter months (Table 2, Year 1). This indicates that the

Table 2
Fruit yield (kg m−2) with or without supplemental intra-canopy LED illumination.
Autumn Winter Spring Total Class 1†

Year 1
CR 1.14 ± 0.10a 3.44 ± 1.08a 5.18 ± 0.59a 9.75 ± 1.09a 8.51 ± 1.24a
LED* 0.82 ± 0.25a 3.48 ± 0.46a 7.03 ± 0.84b 11.32 ± 0.95a 10.12 ± 0.87a
WW 0.45 3.82 6.54* 10.81 9.95
CW 0.90 3.07 7.99** 11.96 10.72
RB 1.00 3.10 6.14 10.24 8.96
RGB 0.91 3.92 7.43* 12.26 10.85
Year 2
CR 2.36 ± 0.59a 2.76 ± 0.22a 4.99 ± 0.39a 10.11 ± 0.86a 9.37 ± 0.86a
LED 2.94 ± 0.30a 3.20 ± 0.30a 6.45 ± 0.73b 12.60 ± 1.23b 11.81 ± 1.06b

Yield shown for autumn (Nov.-Dec.), winter (Jan.-Feb.) and spring (Mar.-Apr.) represents the total obtained yield. ‘Class 1′ is the fraction of high, export-quality fruit
yield from the ‘Total’ yearly yield. Data represent means ± SD. Year 1: CR – mean of 3 control, non-illuminated plots; LED* – mean of the 4 illuminated plots (WW,
CW, RB, RGB). The yield in each single illuminated plot is also shown. WW, warm white; CW, cool white; RB, red-blue; RGB, red-green-blue. Year 2: CR – mean of 6
non-illuminated plots; LED – mean of 3 plots illuminated with CW. Distinct letters (separately for each year and season) denote statistically significant differences
(p < 0.05). Significant differences between the spring yield in the single LED plots as compared with the average control (year 1), as determined by one-sample t-
test, are denoted by *(p < 0.05) or **(p < 0.01). † The difference between the total fruit yield and class 1 yield represents class 2 fruit yield.

85
N.C. Joshi, et al.
hand, the yield by weight in the LED-illuminated plots was on average
36% higher (Table 2, Year 1 - Spring). The spring yield in each of the
single LED plots was compared to the average CR using one-sample t-
test. This indicated that the differences were significant for WW, CW
and RGB. As the CW LED resulted in the highest yield (> 50% addition
compared to the control) in the spring, it was selected for the second
experiment. The difference in total yield in the first experiment (˜16%
more in LED) was, however, not significant.
Like the first experiment, in the second experiment, in which CW-
LED illumination was replicated in 3 plots, there were no differences
between LED and CR in the autumn and winter. Notably, the increase in
yield in the spring months in year 2 was very similar to year 1, aver-
aging at 29% (Table 2, Year 2 - Spring). In this (second) season, the
overall yield was also significantly higher than the control, resulting in
the addition of ˜25%. This is also reflected in the significant increase, of
26%, obtained in the class 1 (export-quality) yield of LED-illuminated
plots as compared to the control. The difference between the two ex-
periments with regard to the total seasonal yield may have resulted
from the use of a higher light intensity in year 2. In both experiments,
class 1 yield represented a major fraction of the total yield, with no
major differences in this fraction between control and illuminated plots
(of the same experiment). In the first year, class 1 yield amounted to
˜88% of the total yield, and in the second year to ˜93% of the total yield,
in both control and LED (Table 2, ‘Class 1′).
Class 1 yield, whose fraction out of the total yield did not change
between CR and LED, is also shown as fruit number m−2 (Table 3).
Inspection of the data shows the same trends as total kg m−2. The
average added yield in the spring in LED were 45% and 33%, for the
first and second experiment, respectively. These differences were
somewhat higher than the total added yield (by weight) obtained in
LED-illuminated plots, raising the question of whether the addition of
fruit in illuminated plots was accompanied by a decrease in fruit size or
weight compared to fruit of control plots. We thus compared the size
and average fruit weight in control vs. LED. This was conducted for the
spring harvest, when the differences in yield (both by weight and by
fruit number) were obtained. No significant changes in the size or
weight of the fruit between control and illuminated plants were found.
Fruit distribution to size categories (by diameter) was generally the
same in the control and LED-illuminated sections in both experiments
(Fig. 4A and C). Note that the distribution pattern of the spring fruit
into size categories differed between the two experiments, with the
second season fruit having a higher fraction of larger fruit and nearly no
86
Scientia Horticulturae 250 (2019) 81–88
‘small’ category fruit. However, the same pattern was observed for
control and LED in each of the experiments. The average fruit weight in
each size category was likewise unaltered by the illumination in both of
the experiments (Fig. 4B and D).
4. Discussion
Crops grown in regions with ample sunlight year-round are typically
not considered to be light-limited. However, within the canopies of
many crop plants, light can be a limiting factor mostly due to canopy
geometry and hence self-shading, but also because of shading imposed
by neighboring plants, or the geographical positioning (i.e., less light in
the northern side). The light limitation, uneven distribution of light, as
well as differences in the spectral composition exhibited by different
parts of the canopy lead to a reduction in photosynthetic capacity
which may have an overall disadvantage for plant productivity and
fruit yield (Yamori, 2016; Yamori et al., 2016). In this study, we de-
monstrate that supplemental intra-canopy LED illumination can be
utilized as a means to increase the yield of bell peppers grown in passive
tunnels in the Jordan Valley, which exhibits relatively high solar irra-
diance. In this growth regime, temperatures during the season vary,
reflecting the ambient ones. These conditions differ substantially from
the ones employed in other LED-interlighting studies, typically con-
ducted in northern latitude regions in the context of actively-controlled
greenhouses and additional top lighting. Thus, our findings raise the
prospect for utilizing LED-interlighting in regions not necessarily light-
limited (at least not at canopy top) for increasing fruit yield.
The reported effects of intra-canopy illumination for intensively-
grown vegetables (in the context of actively-controlled greenhouses)
vary from case to case. In cucumber, while significantly increasing
photosynthesis in lower canopy leaves, the LED-interlighting did not
result in a significant increase in fruit yield (Trouwborst et al., 2010). In
another study, of mini-cucumber, some fruit yield increase was ob-
tained, though only during part of the growth season (Hao et al., 2012).
In tomato, LED-interlighting resulted in yield increases attributed to the
increase in fruit size and/or number by 24% to 36% (Gómez and
Mitchell, 2016; Pepin et al., 2014). In pepper, it has been shown quite
early on that interlighting, at the time implemented using HPS lamps,
resulted in a 23% increase in total fruit yield (Hovi-Pekkanen et al.,
2006). In later studies, the application of LED-interlighting for sweet
pepper was reported to increase the yield by ˜16%, attributed to higher
fruit number (Guo et al., 2016; Jokinen et al., 2012). In our study, we
Fig. 4. LED intra-canopy illumination does
not affect fruit size and weight. Fruit size
distribution (A and C) and fruit weight (B and
D) in control (CR) and LED-illuminated (LED)
plots in the experiments of seasons 2016–2017
(A and B) and 2017–2018 (C and D). Data are
shown for fruit in each size category, sorted by
diameter: ‘S’, small (< 60 mm); ‘M’, medium
(60–70 mm); ‘L’, large (71–85 mm); ‘XL’, extra-
large (86–95 mm); ‘G’, giant (96–110 mm). The
analysis was carried out on class 1 (export-
quality) fruit harvested in spring months (Mar.-
Apr.), when a significant difference in yield,
attributed to higher number of fruits, was ob-
tained with the added illumination (see
Table 3). Bars denote means ± SD (year 1:
n = 75–401 fruits; year 2: n = 49–328 fruits).
In both experiments, no significant differences
were found between CR and LED.
N.C. Joshi, et al.
found that LED light supplementation significantly enhanced photo-
synthetic rates in inner canopy leaves compared to leaves of control
plants, and resulted in an addition of 25% to the total yield (year 2
experiment). In the growth regime within passive tunnels, this addition
amounted to ˜30% of the yield in the spring months, during which
about half of the total yield is generally obtained. The increase in
productivity was realized by a higher number of fruit, and without
significant changes in fruit size or weight. As shown previously
(Jokinen et al., 2012), we believe that the higher numbers of fruit re-
sulted from higher fruit-set and/or lower abortion rates achieved with
the supplemental illumination.
The added fruit yield in our study was not accompanied by any
change in the fraction of class 1 (export-quality) fruit. This was true for
the first year experiment, when both narrow- and broad-band LEDs
were used, as well as for the second year experiment, employing CW
LEDs. Similar observations were made in other interlighting studies of
pepper (Guo et al., 2016; Jokinen et al., 2012). In tomato, it was shown
that the application of narrow-band red and blue light as intra-canopy
illumination did not pose any negative effects on fruit quality, as as-
sessed both by sensory panels and physicochemical parameters
(Dzakovich et al., 2015). Furthermore, Guo et al. found increases in the
content of health-promoting compounds in pepper fruits originating
from plots illuminated by LED-interlighting as opposed to HPS top
lighting (Guo et al., 2016).
LED-interlighting products most commonly consist of blue- and red-
LED chip combinations, specifically targeted for excitation of the
chlorophyll pigments and thus for enhancing photosynthetic activity
(e.g., ‘GreenPower LED interlighting module’, Philips Lighting Holding
B.V., www.philips.com/horti). Nonetheless, additional spectral com-
positions, including different ratios of blue, red, far-red and white light
have been tested for interlighting (Gómez and Mitchell, 2016; Hao
et al., 2012; Hernández and Kubota, 2012; Lu et al., 2012). Here we
showed that cool-white light, with dominant green and blue light
components (Fig. 2, CW), resulted in an overall 25% increase in bell
pepper fruit yield. Beneficial LED-interlighting utilizing solely white
LEDs has also been demonstrated for tomato (Lu et al., 2012), where it
resulted in a yield increase of 12%, as compared to 14% using red light.
Application of a single spectral LED component might be preferential to
a combination of LEDs in linear fixtures, especially when used within
dense canopies, as the light spectra exhibited by the nearest sur-
rounding foliage would be more homogeneous.
In our first experiment, the highest yields were obtained with the
CW and RGB LEDs (Table 2). In fact, the lowest yield out of all LED
variants was obtained with RB - this being the typical LED combination
utilized for interlighting. Although these were not repeated in the
second experiment, the results raise the possibility that the green
spectral component, whether broad- or narrow-band, and which is
dominant in cool-white, may be advantageous for use in intra-canopy
illumination. One reasoning for this may be the improved penetration
of green- as opposed to blue- or red light into the canopy (Lu et al.,
2012; Smith et al., 2017). Also supporting this is the finding that Yellow
photoselective nets, which enhance the green part of the spectrum,
resulted in increased productivity in sweet pepper (Shahak et al., 2016).
This direction, in relation to the spectra of intra-canopy illumination,
necessitates further investigation.
In addition to spectral composition, a multitude of other parameters
related to the practical application of interlighting may affect the im-
pacts of the illumination on crop productivity and yield. These include
the duration of the illumination, timing (i.e., day vs. night or summer vs.
winter; Li et al., 2016; Tewolde et al., 2016), intensity, positioning
within the canopy and direction of the light (Song et al., 2016). Thus,
achieving the desired practical effects of intra-canopy illumination
likely requires testing numerous conditions, in addition to specific ad-
justments for different crop species/cultivars and the growth condi-
tions. In the case of passive tunnels in our region, light supplementation
starting in Nov. resulted in increased yields in Mar. and Apr. It remains
87
Scientia Horticulturae 250 (2019) 81–88
to be determined whether providing the illumination earlier in the
season can also increase the yields obtained in winter months.
In sum, in this study we showed that LED interlighting can be uti-
lized for increasing the yield of bell pepper at conditions not previously
tested, namely within passive tunnels in which canopy top is generally
not light-limited. The ability to obtain increased yield in the spring
months is economically beneficial for the Jordan Valley region.
However, further experimentation for optimization of the LED infra-
structure, as well as the illumination regime, is still essential in order to
efficiently apply intra-canopy illumination under the local growth re-
gime. The use of intra-canopy illumination in such conditions may
become relevant especially if or when coupled to photovoltaic systems
in the future, concomitant with the ongoing improvements in LEDs
efficiency along with the decline in their prices.
Acknowledgements
We thank Merav Hajaj and Yuval Bussi (Weizmann Institute of
Science), Marc Perel (Min. of Agriculture) and Assa Florentin (ARO) for
their assistance with data collection and analysis. We also wish to thank
David Silverman, Tamar Alon (Agricultural Extension Service), Uri
Adler, Itzhak Esquira (Plants Board) and Hagai Yasuor (ARO) for the
fruitful discussions. This work was supported by the Israeli Plants
Production and Marketing Board and the Jewish National Fund (KKL).
N.C.J. was supported in part by an ARO Postdoctoral Fellowship – India
and China. This work is dedicated to the memory of Naftali Zur, who
passed away in early 2018.
Appendix A. Supplementary data
Supplementary material related to this article can be found, in the
online version, at doi:https://doi.org/10.1016/j.scienta.2019.02.039.
References
Aloni, B., Karni, L., Zaidman, Z., Schaffer, A.A., 1996. Changes of carbohydrates in pepper
(Capsicum annuum L.) flowers in relation to their abscission under different shading
regimes. Ann. Bot. 78, 163–168. https://doi.org/10.1006/anbo.1996.0109.
Aloni, B., Karni, L., Zaidman, Z., Schaffer, A.A., 1997. The relationship between sucrose
supply, sucrose-cleaving enzymes and flower abortion in pepper. Ann. Bot. 79,
601–605. https://doi.org/10.1006/anbo.1996.0410.
Bantis, F., Smirnakou, S., Ouzounis, T., Koukounaras, A., Ntagkas, N., Radoglou, K., 2018.
Current status and recent achievements in the field of horticulture with the use of
light-emitting diodes (LEDs). Sci. Hortic. (Amsterdam) 235, 437–451. https://doi.
org/10.1016/j.scienta.2018.02.058.
Carvalho, S.D., Schwieterman, M.L., Abrahan, C.E., Colquhoun, T.A., Folta, K.M., 2016.
Light quality dependent changes in morphology, antioxidant capacity, and volatile
production in sweet basil (Ocimum basilicum). Front. Plant Sci. 7, 1–14. https://doi.
org/10.3389/fpls.2016.01328.
Chen, Xli, Xue, X., zhang, Guo, zhong, W., Wang, Lchun, Qiao, Xjun, 2016. Growth and
nutritional properties of lettuce affected by mixed irradiation of white and supple-
mental light provided by light-emitting diode. Sci. Hortic. (Amsterdam) 200,
111–118. https://doi.org/10.1016/j.scienta.2016.01.007.
D’Souza, C., Yuk, H.G., Khoo, G.H., Zhou, W., 2015. Application of light-emitting diodes
in food production, postharvest preservation, and microbiological food safety.
Compr. Rev. Food Sci. Food Saf. 14, 719–740. https://doi.org/10.1111/1541-4337.
12155.
Demers, D.A., Dorais, M., Wien, C.H., Gosselin, A., 1998. Effects of supplemental light
duration on greenhouse tomato (Lycopersicon esculentum Mill.) plants and fruit
yields. Sci. Hortic. (Amsterdam) 74, 295–306. https://doi.org/10.1016/S0304-
4238(98)00097-1.
Dorais, M., 2003. The use of supplemental lighting for vegetable crop production: light
intensity, crop response, nutrition, crop management, cultural practices. Can.
Greenh. Conf. 1–8. https://doi.org/10.1017/CBO9781107415324.004.
Dueck, T.A., Grashoff, C., Broekhuijsen, G., Marcelis, L.F.M., 2006. Efficiency of light
energy used by leaves situated in different levels of a sweet pepper canopy. Acta
Hortic. 711, 201–205. https://doi.org/10.17660/ActaHortic.2006.711.25.
Dzakovich, M.P., Gómez, C., Mitchell, C.A., 2015. Tomatoes grown with light-emitting
diodes or high-pressure sodium supplemental lights have similar fruit-quality attri-
butes. HortScience 50, 1498–1502.
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.
Goldberg, B., Klein, W.H., 1977. Variations in the spectral distribution of daylight at
various geographical locations on the earth’s surface. Sol. Energy 19, 3–13. https://
doi.org/10.1016/0038-092X(77)90083-4.
N.C. Joshi, et al.
Gómez, C., Mitchell, C.A., 2014. Supplemental lighting for greenhouse-grown tomatoes:
intracanopy LED Towers vs. Overhead HPS lamps. Acta Hortic. 1037, 855–862.
Gómez, C., Mitchell, C.A., 2016. In search of an optimized supplemental lighting spec-
trum for greenhouse tomato production with intracanopy lighting. Acta Hortic. 1134,
57–62. https://doi.org/10.17660/ActaHortic.2016.1134.8.
Gómez, C., Morrow, R.C., Bourget, C.M., Massa, G.D., Mitchell, C.A., 2013. Comparison of
intracanopy light-emitting diode towers and overhead high-pressure sodium lamps
for supplemental lighting of greenhouse-grown tomatoes. Horttechnology 23, 93–98.
https://doi.org/10.1017/CBO9781107415324.004.
González-Real, M.M., Liu, H.Q., Baille, A., 2009. Influence of fruit sink strength on the
distribution of leaf photosynthetic traits in fruit-bearing shoots of pepper plants
(Capsicum annuum L.). Environ. Exp. Bot. 66, 195–202. https://doi.org/10.1016/j.
envexpbot.2009.01.005.
Gunnlaugsson, B., Adalsteinsson, S., 2006. Interlight and plant density in year-round
production of tomato at northern latitudes. Acta Hortic. 711, 71–75. https://doi.org/
10.17660/ActaHortic.2006.711.6.
Guo, X., Hao, X., Khosla, S., Kumar, K.G.S., Cao, R., Bennett, N., 2016. Effect of LED
interlighting combined with overhead HPS light on fruit yield and quality of year-
round sweet pepper in commercial greenhouse. Acta Hortic. 1134, 71–78. https://
doi.org/10.17660/ActaHortic.2016.1134.10.
Hao, X., Zheng, J., Little, C., Khosla, S., 2012. LED inter-lighting in year-round green-
house mini-cucumber production. Acta Hortic. 956, 335–340.
Hemming, S., 2011. Use of natural and artificial light in horticulture -interaction of plant
and technology. Proceedings of the VI International Symposium on Light in
Horticulture 15–19. https://doi.org/10.17660/ActaHortic.2011.907.1.
Hernández, R., Kubota, C., 2012. Tomato seedling growth and morphological responses to
supplemental LED lighting red:blue ratios under varied daily solar light integrals.
Acta Hortic. 956, 187–194. https://doi.org/10.17660/ActaHortic.2012.956.19.
Hovi, T., Näkkilä, J., Tahvonen, R., 2004. Interlighting improves production of year-
round cucumber. Sci. Hortic. (Amsterdam) 102, 283–294. https://doi.org/10.1016/j.
scienta.2004.04.003.
Hovi-Pekkanen, T., Näkkilä, J., Tahvonen, R., 2006. Increasing productivity of sweet
pepper with interlighting. Acta Hortic. 711, 165–170. https://doi.org/10.17660/
ActaHortic.2006.711.19.
Jacovides, C.P., Tymvios, F.S., Asimakopoulos, D.N., Theofilou, K.M., Pashiardes, S.,
2003. Global photosynthetically active radiation and its relationship with global solar
radiation in the Eastern Mediterranean basin. Theor. Appl. Climatol. 74, 227–233.
https://doi.org/10.1007/s00704-002-0685-5.
Jokinen, K., Särkkä, L.E., Näkkilä, J., 2012. Improving sweet pepper productivity by LED
interlighting. Acta Hortic. 956, 59–66.
Korczynski, P.C., Logan, J., Faust, J.E., 2002. Mapping monthly distribution of daily light
integrals across the contiguous United States. Horttechnology 12, 12–16.
Li, Q., Kubota, C., 2009. Effects of supplemental light quality on growth and phyto-
chemicals of baby leaf lettuce. Environ. Exp. Bot. 67, 59–64. https://doi.org/10.
1016/j.envexpbot.2009.06.011.
Li, X., Lu, W., Hu, G., Wang, X.C., Zhang, Y., Sun, G.X., Fang, Z., 2016. Effects of light-
emitting diode supplementary lighting on the winter growth of greenhouse plants in
the yangtze river delta of China. Bot. Stud. 57. https://doi.org/10.1186/s40529-015-
0117-3.
Lu, N., Maruo, T., Johkan, M., Hohjo, M., Tsukagoshi, S., Ito, Y., Ichimura, T., Shinohara,
Y., 2012. Effects of supplemental lighting with light-emitting diodes (LEDs) on to-
mato yield and quality of single-truss tomato plants grown at high planting density.
Environ. Control Biol. https://doi.org/10.2525/ecb.50.63.
Marcelis, L.F.M., Heuvelink, E., Baan Hofman-Eijer, L.R., Den Bakker, J., Xue, L.B., 2004.
Flower and fruit abortion in sweet pepper in relation to source and sink strength. J.
Exp. Bot. 55, 2261–2268. https://doi.org/10.1093/jxb/erh245.
Massa, G.D., Kim, H.H., Wheeler, R.M., Mitchell, C.A., 2008. Plant productivity in re-
sponse to LED lighting. HortScience 43, 1951–1956.
Mitchell, C.A., 2015. Academic research perspective of LEDs for the horticulture industry.
HortScience 50, 1293–1296.
88
Scientia Horticulturae 250 (2019) 81–88
Mitchell, C.A., Dzakovich, M.P., Gomez, C., Lopez, R., Burr, J.F., Hernández, R., Kubota,
C., Currey, C.J., Meng, Q., Runkle, E.S., Bourget, C.M., Morrow, R.C., Both, A.J.,
2015. Light-emitting diodes in horticulture. Hortic. Rev. (Am. Soc. Hortic. Sci). 43,
1–88. https://doi.org/10.1002/9781119107781. ch01.
Morrow, R.C., 2008. LED lighting in horticulture. HortScience 43, 1947–1950.
Nelson, J.A., Bugbee, B., 2014. Economic analysis of greenhouse lighting: light emitting
diodes vs. High intensity discharge fixtures. PLoS One 9. https://doi.org/10.1371/
journal.pone.0099010.
Olle, M., Virsile, A., 2013. The effects of light-emitting diode lighting on greenhouse plant
growth and quality. Agric. Food Sci. 22, 223–234. https://doi.org/10.1016/j.
envexpbot.2009.06.011.
Ouzounis, T., Rosenqvist, E., Ottosen, C.-O., 2015. Spectral effects of artificial light on
plant physiology and secondary metabolism. Hortscience 50, 1128–1135.
Pepin, S., Fortier, E., Béchard-Dubé, S.A., Dorais, M., Ménard, C., Bacon, R., 2014.
Beneficial effects of using a 3-D LED interlighting system for organic greenhouse
tomato grown in Canada under low natural light conditions. Acta Horticulturae.
International Society for Horticultural Science (ISHS) 239–246. https://doi.org/10.
17660/ActaHortic.2014.1041.28.
Shahak, Y., 2008. Photo-selective netting for improved performance of horticultural
crops. A review of ornamental and vegetable studies carried out in Israel. Acta Hortic.
161–168. https://doi.org/10.17660/ActaHortic.2008.770.18.
Shahak, Y., Kong, Y., Ratner, K., 2016. The wonders of yellow netting. Acta Hortic. 1134,
327–334. https://doi.org/10.17660/ActaHortic.2016.1134.43.
Shifriss, C., Pilowsky, M., Aloni, B., 1994. Variation in flower abscission of peppers under
stress shading conditions. Euphytica 78, 133–136. https://doi.org/10.1007/
BF00021408.
Singh, D., Basu, C., Meinhardt-Wollweber, M., Roth, B., 2015. LEDs for energy efficient
greenhouse lighting. Renew. Sustain. Energy Rev. 49, 139–147. https://doi.org/10.
1016/j.rser.2015.04.117.
Smith, H.L., McAusland, L., Murchie, E.H., 2017. Don’t ignore the green light: exploring
diverse roles in plant processes. J. Exp. Bot. 68, 2099–2110. https://doi.org/10.
1093/jxb/erx098.
Song, Y., Jiang, C., Gao, L., 2016. Polychromatic supplemental lighting from underneath
canopy is more effective to enhance tomato plant development by improving leaf
photosynthesis and stomatal regulation. Front. Plant Sci. 7, 1832. https://doi.org/10.
3389/fpls.2016.01832.
Tewolde, F.T., Lu, N., Shiina, K., Maruo, T., Takagaki, M., Kozai, T., Yamori, W., 2016.
Nighttime supplemental LED inter-lighting improves growth and yield of single-truss
tomatoes by enhancing photosynthesis in both winter and summer. Front. Plant Sci.
7, 448. https://doi.org/10.3389/fpls.2016.00448.
Thimijan, R.W., Heins, R.D., 1983. Photometric, radiometric, and quantum light units of
measure: a review of procedures for interconversion. HortScience 18, 818–822.
Trouwborst, G., Oosterkamp, J., Hogewoning, S.W., Harbinson, J., van Ieperen, W., 2010.
The responses of light interception, photosynthesis and fruit yield of cucumber to
LED-lighting within the canopy. Physiol. Plant. 138, 289–300. https://doi.org/10.
1111/j.1399-3054.2009.01333.x.
Turner, A.D., Wien, H.C., 1994. Dry matter assimilation and partitioning in pepper cul-
tivars differing in susceptibility to stress-induced bud and flower abscission. Ann. Bot.
https://doi.org/10.1006/anbo.1994.1077.
Wubs, A.M., Heuvelink, E., Marcelis, L.F.M., 2009. Abortion of reproductive organs in
sweet pepper (Capsicum annuum L.): a review. J. Hortic. Sci. Biotechnol. 84, 467–475.
https://doi.org/10.1080/14620316.2009.11512550.
Yamori, W., 2016. Photosynthetic response to fluctuating environments and photo-
protective strategies under abiotic stress. J. Plant Res. 129, 379–395. https://doi.org/
10.1007/s10265-016-0816-1.
Yamori, W., Kondo, E., Sugiura, D., Terashima, I., Suzuki, Y., Makino, A., 2016. Enhanced
leaf photosynthesis as a target to increase grain yield: insights from transgenic rice
lines with variable Rieske FeS protein content in the cytochrome b6/f complex. Plant
Cell Environ. 39, 80–87. https://doi.org/10.1111/pce.12594.