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Acta Hortic. 1296. ISHS 2020. DOI 10.17660/ActaHortic.2020.1296.104
Proc. Int. Symp. on Advanced Technologies and Management for
Innovative Greenhouses – GreenSys2019
Eds.: P.E. Bournet et al.
823
Evaluation of four soilless substrate systems for
greenhouse strawberry production
T. McKean1, a, M. Kroggel1, C. Kubota1 and R. Naasz2
1Department of Horticulture and Crop Science, The Ohio State University, Columbus, Ohio, USA; 2PremierTech
Horticulture, 1 Avenue Premier, Riviere-du-Loup, Québec, Canada.
Abstract
Growing demand for locally produced strawberries (Fragaria × ananassa) with
high flavor quality has led to an increase in US soilless strawberry production under
controlled environment in recent years. Soilless substrate-based production systems
need to be optimized for plant productivity, plant health, and ultimately grower profit.
This study investigated effects of four different substrate systems, which included two
commercial substrate mixes, on ‘Albion’ plant vigor, fruit yield, and fruit quality. The
first commercial mixture (Com1) was used with a plastic trough (container) and
contained peat moss, perlite, coco coir chips, supplemental arbuscular mycorrhizal
fungi (Glomus intraradices, strain PTB297 syn. Rhizophagus irregularis DAOM 197198)
(AMF), and biofungicide bacteria (Bacillus pumilus, strain PTB180) (BP) while the
second commercial mixture (Com2) was composed of peat moss, composted pine bark,
coco coir fiber, AMF, and BP, packaged in a plastic bag. Our standard lab substrate
mixture (StMix-S) for strawberry production consisted of perlite, fine particle coco coir,
and peat moss. Additionally, this same mix was used in the same plastic trough but
supplemented with AMF (StMix-M). Forty-eight ‘Albion’ plants were grown in each of
the four systems from October 2017 to May 2018 in a north-south oriented glass
greenhouse (Columbus, OH, USA). Com1 and Com2 had 26-31% greater cumulative
fruit yield per plant (619.9±35.6 to 652.5±34.1 g) than StMix-S (476.3±41.0 g) but did
not have significantly different yields than StMix-M (575.6±32.7 g). AMF colonization
was confirmed in StMix-M but not in Com1 nor Com2. BP was confirmed in Com1 and
Com2. These findings suggest that physical, chemical, and biological properties must
be integrated in order to maximize soilless strawberry yield.
Keywords: Fragaria × ananassa, hydroponics, greenhouse strawberry, soilless culture,
arbuscular mycorrhizal fungi, plant growth-promoting rhizobacteria
INTRODUCTION
The United States strawberry (Fragaria × ananassa) industry is dominated by California
open-field production (~90%) (Samtani et al., 2019). Soil-based production systems face
numerous challenges including high soilborne pathogen pressure and limited availability of
effective soil fumigants. Additionally, demand for a year-round supply of high-quality
strawberries is increasing in North America, and the current supply chain involves long
distance transportation, which can reduce produce quality. Soilless substrate-based
greenhouse production systems alleviate these challenges by starting with new growing
media (substrate) in each production cycle and by extending the production season by
growing in a controlled environment. Greenhouse soilless production of strawberries is more
common in parts of Europe (Lieten, 2005) but has not been widely adopted in North America
(with only ~16 ha in Canada) (Samtani et al., 2019). Several studies have investigated
controlled environment soilless production in North America and have proven that it is a
potential alternative to open-field production especially for localized off-season (November-
March) production in non-optimal environments (Cantliffe et al., 2007; Kempler, 2004;
Paranjpe and Cantliffe, 2003; Paranjpe et al., 2008). One way to increase the viability of this
new production method is to optimize soilless strawberry substrate systems for plant
aE-mail: mckean.50@osu.edu
824
productivity, plant health, and grower profit.
In soilless substrate systems, desired root-zone physical and chemical characteristics
are often achieved by mixing selected organic or inorganic substrate components. Container
selection is also important as different container geometries impact the effects of intrinsic
substrate properties. Thus, substrate system (combination of container and substrates)
optimization is desired. One important physical property for strawberry substrate systems is
high air porosity to allow gas exchange and adequate drainage (Wallach, 2008). Air porosity
must also be balanced with adequate water holding capacity to ensure water and nutrient
uptake (Wallach, 2008). Substrate system design must also consider optimal root-zone
chemical characteristics. Ideal root-zone pH for substrate food crop production is between 5.5
and 6.5 to keep nutrients in plant available forms (Savvas et al., 2013). Strawberry plants
prefer a relatively low electrical conductivity (EC). For example, an EC between 1.1 and 2.1 dS
m-1 was found to produce the greatest plant growth and yield in seven tested cultivars (Sun et
al., 2015). Comparisons in strawberry plant growth and fruit yield between different
substrates were made by various research groups (e.g., Cantliffe et al., 2007; Depardieu et al.,
2016; Kuisma et al., 2014; Recamales et al., 2007). During the past several years, we have
advised growers to use a mixture of perlite, coconut coir, and peat moss (2:1:1 volume ratio)
to meet the optimum root-zone properties (Kubota and Kroggel, 2008, unpublished data).
However, mixing substrate components is sometimes laborious increasing production costs.
Commercially available, pre-mixed substrates can replace our custom mixed substrates if they
can achieve desirable characteristics, leading to comparable or possibly greater fruit yields
and quality.
Recently, use of microbiological amendments have been commonly practiced in
horticultural substrate mix to encourage growth and prevent disease. Strawberry plants
grown in the open field have a diverse rhizosphere, and these naturally occurring rhizosphere
microbes protect plants from abiotic and biotic stresses. Inoculation of substrates with plant
growth-promoting rhizobacteria (PGPR) and arbuscular mycorrhizal fungi (AMF) has been
proposed to increase strawberry plant performance in soilless production systems (Vestberg
et al., 2004). PGPR provide many ecosystem services including nitrogen fixation, competing
with pathogenic microbes, and synthesizing various plant hormones (Bhattacharyya and Jha,
2012). AMF provides similar observable benefits to colonized plants but is primarily known
to increase water and nutrient uptake by creating a larger effective root surface area with its
hyphae extensions (Smith and Smith, 2011). The objective of the present study is to compare
plant growth, fruit yield, and fruit quality of strawberry plants grown in two commercial
substrate mixture-based systems, and our standard system with and without AMF.
MATERIALS AND METHODS
Greenhouse and growing conditions
The experiment was conducted in a 103 m2 growing area in a glass greenhouse (40°N;
83°W) with a concrete floor and a north-south ridge orientation. The growing area comprised
six rows of metal gutters (5.2×0.3 m) (Meteor Systems, Breda, The Netherlands) oriented
north-south at a height of 0.9 m above the floor and spaced 1 m center to center. Supplemental
LED lighting (SpecGrade LED, Columbus, OH, USA) was used to provide an average
photosynthetic photon flux density (PPFD) of 200.6±17.5 µmol m-2 s-1 at the plant canopy
level. The lighting was set to turn on 16 h before sunset and to turn off 30 min prior to sunset.
Within the supplemental lighting period, lights were turned off when the exterior solar
radiation reached 400 W m-2 with a 15-min delay to prevent excessive cycling. A 25% shade
(Kool Ray, Continental Products, Euclid, OH, USA) was applied on April 23, 2018 and stayed
on until the final vegetative measurements were made (May 1, 2018).
Greenhouse temperature was controlled with a pad-and-fan evaporative cooling and a
hot-water radiant heating installed at the perimeters of greenhouse. Heating/cooling
setpoints were 20/24°C during daylight, 9/12°C during the night, and 18/24°C when LEDs
lighting was the sole source (before sunrise). Nighttime fogging with under-gutter misters
(Netafim USA, Fresno, CA, USA) was implemented as described in Kroggel and Kubota (2017).
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Air temperature and relative humidity were measured with a fan-aspirated HMP60 probe
(Vaisala, Helsinki, Finland) at the central location of the greenhouse. PPFD was measured at
the plant canopy and in the greenhouse attic using LI-190R quantum sensors (LI-COR, Lincoln,
NE, USA). Additionally, canopy-level air temperature was monitored in each block using
thermocouples (Type T, gauge 24, Omega Inc., Stamford, CT, USA). All sensors readings were
recorded with a CR23X datalogger (Campbell Scientific, Logan, UT, USA).
Throughout the experiment, all plants were drip-irrigated with nutrient solution
(Yamazaki strawberry formula) composed of (mg L-1): 70 NO3-N, 7 NH4-N, 15 PO4-P, 117 K, 40
Ca,16 SO4-S, 12 Mg, and micronutrients. Irrigation runs were 1-min long each distributing 33
mL per run per plant with 6-13 runs a day to achieve a target discharge (efflux solution)
percentage between 10 and 20%. Extra emitters were added as needed to compensate the
difference between substrate systems. Additionally, target ranges of efflux solution pH and
electrical conductivity (EC) were 5.5-6.5 and 1.2-1.8 dS m-1, respectively. Necessary
adjustments to pH and EC were made throughout the experiment based on the individual
substrates used.
Planting materials and substrate systems
‘Albion’ plug plants (McNitt Growers, Carbondale, Illinois, USA) were received on
September 7, 2017, and 75 plants were planted into each substrate in 350 mL tree-band pots
on September 12. On October 23, 48 plants from each substrate were selected for transplant
into the final production system. We used two commercial substrate mixtures (Premier Tech
Horticulture, Riviere-du-Loup, QC, Canada) and two custom substrate mixes based on our
previous laboratory standard protocols. The first commercial substrate (Com1) contained
peat moss, perlite, coco coir chips, AMF (Glomus intraradices, strain PTB297 syn. Rhizophagus
irregularis DAOM 197198), and bio-fungicide bacteria (Bacillus pumilus, strain PTB180) (BP).
The second commercial substrate (Com2) was composed of peat moss, composted pine bark,
coco coir fiber, AMF, and BP. Both commercial mixtures also contained crushed lime and a
starter nutrient mixture. The standard substrate mixture (StMix-S) consisted of a 3:2:1
volumetric ratio of perlite, fine particle coco coir, and peat moss. The fourth substrate was
supplemented with granular AMF (Premier Tech Horticulture, Riviere-du-Loup, QC, Canada)
at 1.5 kg m-3 but otherwise the same as StMix-S. We deviated from our standard 2:1:1 ratio as
the peat moss used was highly acidic. Physical and chemical properties of the substrate
mixtures are summarized in Table 1. Black plastic troughs (99×18×16 cm, Bato Plastics,
Zevenbergen, The Netherlands) were filled with either Com1, StMix-S, or StMix-M and placed
on the gutter. Com2 was a prepacked bag (100×22×8 cm), with drain holes, placed directly on
the gut t e r.
Table 1. Comparison of substrate system properties and components.
Component/Measurement
Com1
Com2
StMix-S
StMix-M
Initial pHa
5.8-6.2
7.0-7.9
4.9-5.0
4.9-5.0
Initial EC (dS m-1)a
0.3
0.7
0.3-0.4
0.3-0.4
Air porosity
22%b
45%b
15%b,
15%b
Water-holding capacity
72%b
47%b
61%b
61%b
Microbial amendments
AMF, BP
AMF, BP
None
AMF
Container
Plastic Trough
Plastic Bag
Plastic Trough
Plastic Trough
Coconut coir
Yes
Yes
Yes
Yes
Peat moss
Yes
Yes
Yes
Yes
Perlite
Yes
No
Yes
Yes
Pine bark
No
Yes
No
No
a Measured after soaking 1:1 ratio of greenhouse source water (municipal water) and substrate overnight.
b Measured at Premier Tech Horticulture using a procedure described by Caron et al. (2007) using a standardized column height (10
cm) and substrate volume (1 L).
826
Experimental design and statistical analysis
The experiment was setup as a randomized complete block design with four substrate
system treatments. The six rows of metal gutters were split into north and south sections to
give a total of 12 gutter sections with each gutter section representing one experimental unit.
The experiment was blocked every two rows from east to west to produce three blocks (four
experimental units per block), and the four substrate system treatments were randomized
within each block with augmentation so that each treatment had approximately equal average
daily light integrals. Each experimental unit contained 16 plants (two troughs or slabs).
ANOVA-protected Tukey’s HSD test (R version 3.5.1) was used for mean separations.
Additionally, a Welch Two Sample t-test (R version 3.5.1) was used for pairing comparisons.
Measurements and data collection
The first fruit was harvested on December 1, and fruit harvest continued to April 30,
2018. Individual fruit were weighed and then classified by marketability. Unmarketable fruit
consisted of underweight berries (<8 g), fruits with erroneous pollination, uneven ripening
during development, and fruit damaged by pests or fungal disease. Approximately once a
month, a cumulative amount (~120 g) of marketable fruit was collected from each replication
and stored at -80°C. For quality analysis, the frozen fruit samples were thawed in a refrigerator
for 12 h. Homogenized samples were centrifuged to obtain supernatant for quantifying total
soluble solid concentration (TSS, °Brix measured by a refractometer, PR-32α, Atago USA,
Bellevue, WA) and titratable acidity (TA). TA was quantified by titrating a 10 mL aliquot of the
supernatant with 0.1 N NaOH to a pH of 8.2. Based on the volume of 0.1 N NaOH added,
titratable acidity in the form of citric acid (C6H8O7) equivalents (g L-1) was calculated following
the procedure found in Sadler and Murphy (2010). A total of four fruit quality sets were
collected and tested during the production cycle. The four fruit collection periods were
December 17, 2017-January 4, 2018 (QS1), January 7-January 17, 2018 (QS2), April 4-April
12, 2018 (QS3), and April 24-April 29, 2018 (QS4). Plant growth measurements were
performed weekly, including longest petiole length (cm), number of leaves (>5 cm leaf blade),
and number of runners. At the end of the experiment (May 14, 2018), number of crowns was
recorded. Substrate samples were sent to the laboratory at Premier Tech Horticulture for
microbial colonization (AMF and BP) analysis.
RESULTS AND DISCUSSION
The present study compared two commercial substrate systems with our two standard
substrate systems. This experiment was intended to compare substrate systems rather than
specific components. Therefore, the substrate systems we found to be optimal may not be the
same under different growing or climate conditions.
Greenhouse environment and substrate conditions
The average air temperature was 21.9±1.9°C during daytime, 14.2±3.2°C during
nighttime, and 19.1±2.7°C when LED lighting was the sole source. The average DLI throughout
the experiment was 17.7±7.1 mol m-2 d-1 (maximum 37.1 mol m-2 d-1). Average drainage efflux
percentage across the production period (Table 2) was higher than our optimal range (10-
20%) due to increased irrigation at the beginning of the cropping cycle when plants were
small (~45% for all systems in the first month after transplant). Efflux pH (Table 2) in Com2
was higher than the optimal range as too much lime was added to the substrate during mixing
(R. Naasz, pers. commun.). Because of the possible iron deficiency in such a high pH range, we
used Fe-EDDHA as the chelated iron source. Efflux pH in the two laboratory systems was low
because the peat used in the mixture was highly acidic. While efflux pH was outside of the
optimal range for each substrate system, no nutrient deficiencies were observed in the crop.
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Table 2. Efflux nutrient solution (% volume relative to influx solution) and its pH and EC as
affected by four substrate systems (October 2017-May 2018).
System
Efflux volume percentage
Efflux pH
Efflux EC (dS m-1)
Com1
24.2±16.4
7.0±0.3
1.5±0.3
Com2
30.5±15.6
7.6±0.2
1.3±0.2
StMix-S
29.4±15.7
4.8±0.6
1.3±0.4
StMix-M
25.8±16.1
5.1±0.6
1.4±0.4
Fruit yield
At the end of production, Com1 and Com2 had 31 and 26% higher total fresh fruit
production (g fresh weight plant-1) than StMix-S (Table 3). While StMix-M had a similar yield
when compared to the other substrate systems, there was a weak trend indicating that AMF
supplementation can increase yields in StMix (p=0.12, Welch Two-Sample t-test). Granular
AMF supplementation was found to significantly increase greenhouse strawberry yield grown
in coconut coir (Robinson-Boyer et al., 2016). When compared on a monthly production basis,
there were no significant fruit production differences between substrate systems in December
2017, February 2018, or March 2018. In January 2018, StMix-M had 22% higher fruit
production per plant than StMix-S. In April 2018, Com1 had a 38-54% higher fruit production
than all other substrate systems. Another point to note is that Com2 was the only system to
have consistent yields above 100 g plant-1 in the four major production months (January-April
2018). Consistently in yield is important as production stability reduces supply uncertainty
along with ensuring consistent labor usage. Additionally, we had a relatively high fruit cull
percentage in the present experiment. Com2 (19.0±0.6%) had a significantly lower fruit cull
percentage than Com1 (23.6±0.6%) and StMix-M (24.1±0.3%). Fruit cull in StMix-S
(21.6±0.8%) was not significantly different than any other system. Reduced fruit cull
increases potential grower revenue and reduces time spent on fruit thinning and sorting
which further increases potential grower profit.
Table 3. Fruit production plant-1 (g fresh weight plant-1 month-1) as affected by four substrate
systems.
System
December
2017
January
2018
February
2018
March
2018
April
2018
Total
Com1
18.6±7.1a
160.6±9.5ab
97.8±3.0a
97.1±20.1a
278.4±9.8a
652.5±34.1a
Com2
24.6±4.2a
160.1±4.7ab
122.0±9.8a
123.6±23.5a
189.5±4.2b
619.9±35.6a
StMix-S
22.2±1.8a
137.1±7.0b
67.1±16.9a
89.5±32.2a
160.4±3.7b
476.3±41.0b
StMix-M
26.4±0.7a
170.5±9.3a
92.5±12.1a
106.4±13.7a
179.9±8.9b
575.6±32.7ab
Mean separation using Tukey’s HSD test (p=0.05). Means with the same letter are not significantly different.
Fruit quality
There was no significant difference in TSS between substrate systems in QS1, QS2, QS4,
or when all four sampling periods were averaged (Table 4). In QS3, TSS in Com1 was 7.1-
10.8% lower than any other substrate system. The average TA across all four sampling periods
was 5.7% lower in Com2 than StMix-M. There was no significant difference in TA between
systems in QS1 or QS2. However, in both QS3 and QS4, Com2 had significantly lower TA than
either of the standard substrate systems. Com1 was not significantly different than any of the
other systems in either QS3 or QS4. Our preliminary analysis of fruit quality sets in similar
substrate systems from both Arizona and Ohio indicated that TA is positively correlated with
temperature (likely fruit temperature) during the fruit development period (data not shown).
The lower TA in Com2 than other systems observed in QS3 and QS4 is most likely due to the
more horizontal growing habit of plants in Com2 (slab bag) which provided more fruit
shading during the day. This unique canopy structure may have effectively lowered the fruit
temperature and thereby TA. Com2 had the most consistent TSS and TA across all four fruit
828
sampling periods. In Com2, TSS was 9.0% or higher in all four sampling periods while TA was
below 9.0 g L-1 in all periods. This equates to a TSS:TA ratio greater than 1 for every sampling
period. These values are favorable for strawberry as the TA range is typically 9.5-11.8 g L-1,
and the TSS range is typically 8-10.1% (Sadler and Murphy, 2010). Additionally, a higher
TSS:TA ratio has been associated with an increased perception of palatability and sweetness
(Jouquand et al., 2008). This ability to maintain increased fruit quality along with increased
yield is a highly desirable trait when selecting optimal substrate systems for greenhouse
strawberry production.
Table 4. Total soluble solid concentration (TSS, measured as Brix) and titratable acidity of
strawberry fruits as affected by four substrate systems.
System
QS1a
QS2
QS3
QS4
Average
TSS (%)
Com1
8.6±0.3 a
8.6±0.5 a
9.1±0.2 b
9.4±0.2 a
8.9±0.2 a
Com2
9.0±0.4 a
9.4±0.2 a
9.8±0.3 a
9.4±0.2 a
9.4±0.2 a
StMix-S
8.7±0.1 a
8.2±0.4 a
10.0±0.1 a
10.5±0.2 a
9.4±0.2 a
StMix-M
8.7±0.2 a
8.7±0.2 a
10.2±0.2 a
10.0±0.2 a
9.4±0.2 a
Titratable acidity (g L-1)
Com1
8.3±0.6 a
8.4±0.2 a
9.7±0.4 ab
9.3±0.2 ab
8.9±0.1 ab
Com2
8.0±0.4 a
8.5±0.5 a
8.9±0.1 b
8.9±0.1 b
8.7±0.2 b
StMix-S
7.8±0.4 a
8.0±0.2 a
10.2±0.4 a
9.8±0.4 a
8.9±0.1 ab
StMix-M
8.2±0.3 a
8.2±0.4 a
10.2±0.2 a
10.0±0.1 a
9.2±0.1 a
a The four fruit collection periods were December 17, 2017-January 4, 2018 (QS1), January 7-January 17, 2018 (QS2), April 4-April
12, 2018 (QS3), and April 24-April 29, 2018 (QS4).
Mean separation using Tukey’s HSD test (p=0.05). Means with the same letter are not significantly different.
Vegetative vigor
Petiole length has been used in the past as an indicator of overall vegetative vigor and
plant performance (Lieten, 2006). At the time of the final vegetative measurement (May 1,
2018), the average longest petiole in Com1 plants was 28.1±0.8 and 26.5±0.3 cm in Com2 with
both of these commercial systems being significantly longer than StMix-S (21.3±0.8 cm) and
StMix-M (21.8±0.9 cm). There was no difference in average final longest petiole length
between StMix-S and StMix-M (p=0.60, Welch Two-Sample t-test) indicating that AMF
colonization did not have a large effect on vegetative vigor. Martinez et al. (2013) also found
that AMF did not have a significant effect on strawberry plant height.
Microbial colonization
BP colonization (measured as colony forming units, CFU, g-1 of substrate) was confirmed
in Com1 (1.0±0.1×105 CFU g-1) and Com2 (7.5±0.2×104 CFU g-1). No AMF root colonization
was found in Com1 or Com2 (AMF presence in 0 of 16 root samples). However, there was AMF
presence in 38% of StMix-M samples (6 of the 16 root samples). The lack of AMF colonization
in both commercial substrates could be due to the increased availability of phosphorous at
planting from the added starter nutrient (5-40 mg L-1 PO4-P, measured from saturated media
extract) in addition to nutrient solution phosphorous (15 mg L-1). Phosphorous has been
shown to negatively affect AMF colonization in plant roots (Nouri et al., 2014) Root staining
to confirm AMF colonization was performed only once, 7 weeks after transplanting, and roots
were not tested for AMF colonization at the end of the experiment. Therefore, it is possible
that colonization could have occurred later in the cropping cycle. Thus, in the present
experiment, it is not known whether any increases in Com1 or Com2 yield, quality, and
vegetative vigor is due to AMF. However, since BP colonization was confirmed in both Com1
and Com2, there is a potential BP biostimulant effect causing the increases in yield, quality,
and vigor. While neither of the commercial systems had successful AMF colonization, a system
combining both AMF and BP could be desirable. Previous studies have shown that combining
both AMF and PGPR is complex. Certain single PGPR or AMF inoculations increased vegetative
829
growth in peat-grown strawberry plants (Vestberg et al., 2004). However, in the same study,
none of the PGPR or AMF inoculums consistently increased growth when combined with other
strains. Similarly, vegetative vigor was inconsistently affected by single and combined
inoculations of AMF and PGPR in Palencia et al. (2015). These findings indicate that AMF and
PGPR strains must be carefully selected and verified to have synergistic effects.
In other strawberry experiments, BP has been suggested to provide up to a 10%
increase in yield when added to substrate (R. Naasz, pers. commun.). If we take the 10%
increase in yield as PGPR contribution, and there is no yield effect attributed to AMF nor any
other factor, we speculate that the physical characteristics of Com1 and Com2 substrate mixes
might have caused around 16-21% of the increased yield when compared to StMix-S. A
physical characteristic leading to the increased yields in Com1 and Com2 might be the high
air porosity (Table 1). Com1 and Com2 substrates had 7 and 32% higher air porosity than
StMix-S, respectively. Strawberry roots have been found to uptake oxygen at higher rates
compared to other greenhouse crops at the same root temperatures (Inden, 1953), suggesting
the sensitivity of strawberry to oxygen availability in the root zone. Therefore, this relatively
high porosity in the commercial substrate systems could have contributed to the increased
yield when compared to StMix-S.
CONCLUSIONS
The present study shows that integrating chemical, physical, and biological properties
into substrate system design was critical to maximize soilless strawberry production.
Additionally, increased yields from commercial substrate mixes or microbial amendments
could potentially be worth any increased growing material costs.
ACKNOWLEDGEMENTS
The authors thank Catherine Viel for her consultation and Justin Clifford for his
assistance in managing the crop. We also acknowledge Premier Tech Horticulture for the
financial support, and Dosatron, Meteor Systems, and SpecGrade LED for providing in-kind
support to this project.
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