Closing The Yield Gap For Cannabis: A Meta-Analysis of Factors Determining Cannabis Yield

REVIEW
published: 24 April 2019

doi: 10.3389/fpls.2019.00495
Closing the Yield Gap for Cannabis: A

Meta-Analysis of Factors
Determining Cannabis Yield
Rachel Backer 1*, Timothy Schwinghamer 1 , Phillip Rosenbaum 2 , Vincent McCarty 2 ,
Samuel Eichhorn Bilodeau 3 , Dongmei Lyu 1 , Md Bulbul Ahmed 2 , George Robinson 4 ,
Mark Lefsrud 3 , Olivia Wilkins 2 and Donald L. Smith 1
1
Crop Physiology Laboratory, Department of Plant Science, McGill University, Sainte-Anne-de-Bellevue, QC, Canada, 2 Plant
Systems Biology Laboratory, Department of Plant Science, McGill University, Sainte-Anne-de-Bellevue, QC, Canada,
3
Biomass Production Laboratory, Department of Bioresource Engineering, McGill University, Sainte-Anne-de-Bellevue, QC,
Canada, 4 Ravenquest Biomed, Inc., Vancouver, BC, Canada
Until recently, the commercial production of Cannabis sativa was restricted to varieties
that yielded high-quality fiber while producing low levels of the psychoactive cannabinoid
tetrahydrocannabinol (THC). In the last few years, a number of jurisdictions have legalized
the production of medical and/or recreational cannabis with higher levels of THC, and
other jurisdictions seem poised to follow suit. Consequently, demand for industrial-scale
Edited by: production of high yield cannabis with consistent cannabinoid profiles is expected to
Luis A. N. Aguirrezabal,
increase. In this paper we highlight that currently, projected annual production of cannabis
National University of Mar del Plata,
Argentina is based largely on facility size, not yield per square meter. This meta-analysis of cannabis
Reviewed by: yields reported in scientific literature aimed to identify the main factors contributing to
Debora Nercessian, cannabis yield per plant, per square meter, and per W of lighting electricity. In line with
CONICET Mar del Plata, Argentina
Robert VanBuren,
previous research we found that variety, plant density, light intensity and fertilization
Michigan State University, influence cannabis yield and cannabinoid content; we also identified pot size, light type
United States
and duration of the flowering period as predictors of yield and THC accumulation.
*Correspondence:
We provide insight into the critical role of light intensity, quality, and photoperiod in
Rachel Backer
[email protected] determining cannabis yields, with particular focus on the potential for light-emitting
diodes (LEDs) to improve growth and reduce energy requirements. We propose that
Specialty section:
the vast amount of genomics data currently available for cannabis can be used to
This article was submitted to
Crop and Product Physiology, better understand the effect of genotype on yield. Finally, we describe diversification
a section of the journal that is likely to emerge in cannabis growing systems and examine the potential role
Frontiers in Plant Science
of plant-growth promoting rhizobacteria (PGPR) for growth promotion, regulation of
Received: 31 October 2018
Accepted: 01 April 2019
cannabinoid biosynthesis, and biocontrol.
Published: 24 April 2019 Keywords: cannabis, genomics, transcriptomics, chemotype, yield gap, light emitting diodes, PGPR, GWAS
Citation:
Backer R, Schwinghamer T,
Rosenbaum P, McCarty V, Eichhorn INTRODUCTION: CHANGING ATTITUDES ON CANNABIS AND
Bilodeau S, Lyu D, Ahmed MB,
Robinson G, Lefsrud M, Wilkins O and
CURRENT KNOWLEDGE GAPS
Smith DL (2019) Closing the Yield Gap
for Cannabis: A Meta-Analysis of Currently cannabis laws are changing rapidly around the world, with legalization of medical use
Factors Determining Cannabis Yield. appearing in many jurisdictions, followed by legalization of recreational use. In Canada, this has
Front. Plant Sci. 10:495. led to significantly lower barriers to obtaining a license to conduct scientific research under the
doi: 10.3389/fpls.2019.00495 newly adopted Cannabis Act, in comparison with the Access to Cannabis for Medical Purposes
Frontiers in Plant Science | www.frontiersin.org 1 April 2019 | Volume 10 | Article 495

Backer et al. Closing the Cannabis Yield Gap
Regulations (ACMPR) and its predecessor acts: Marihuana for

Medical Purposes Regulations (MMPR) and Marihuana Medical
Access Regulations (MMAR) (Canada, 2001, 2013, 2016, 2018).
However, in the United Sates, while recreational cannabis has
been legalized in nine states and medical cannabis has been
legalized in 21 states (http://www.governing.com/gov-data/
safety-justice/state-marijuana-laws-map-medical-recreational.
html), cannabis remains illegal at the federal level, presenting a
major barrier to research. To meet projected demand for medical
and recreational cannabis products, the yield gap must be closed
with the use of modern scientific tools.
Cannabis is one of the oldest cultivated crops and is
used for food (seeds), fiber (stems), and drugs (flowers); it
was domesticated in Central Asia over 6,000 BCE (Li, 1973;
Mercuri et al., 2002; Clarke and Merlin, 2013, 2016). This
genus produces over 200 secondary metabolites, including
terpenes, phenolic acids and cannabinoids (Andre et al., 2016).
In particular, medical and recreational cannabis are cultivated
for, tetrahydrocannabinol (THC), and cannabidiol (CBD), which
produce physiological and intoxicating effects in humans, which
have been associated with both positive and negative health
outcomes (Hill et al., 2012; Giacoppo et al., 2014; Volkow et al.,
2014; Burstein, 2015; Van Amsterdam et al., 2015). Because
cannabis naturally contains THC and CBD, this plant has been
listed as a controlled substance for the last several decades in
jurisdictions worldwide. Restrictions around cultivation of this
plant has led to a void of scientific research.
For cannabis, the yield gap constitutes the difference between
the maximum possible flower yield compared to current yields
obtained in commercial production. In addition, there is the
important consideration of cannabinoid concentration and
profile, which together determine the quality of the product.
Legal cannabis-producing operations in Canada, show projected
yields that range from 3.36 to 3590 g dry flower m−2 (Figure 1,
Table S1) with MedReleaf achieving the highest yields per square
meter. The first question that must be answered is: is this the
physiological maximum of cannabis plants? The second question
is which production conditions lead to obtaining these high
yields? Another point requiring clarification is whether the most
important yield is in fact the dry flowers (which contain the
highest concentration of medicinal compounds) or the whole
plant (for extraction of medicinal compounds, even from stems
and leaves, which contain significantly lower concentrations).
To date, a limited number of studies have examined factors
contributing to the cannabis yield gap. First, a body of literature
has developed to provide a detailed knowledge base about FIGURE 1 | Cannabis production in Canada; facilities are numbered by facility
existing cannabis strains, at the molecular level. Studies have size (A). Annual production tends to increase with facility size (B) not yield per
square meter (C). It is important to note that it is unclear if facility size is always
begun to elucidate the genetic structure and diversity of cannabis equal to the area of the cannabis production space. Blue dots are projected
(Sawler et al., 2015; Welling et al., 2016a), understand the yields; orange stars are actual yields and correspond to AB Labs (Facility #5),
inheritance of chemotype (De Meijer et al., 2003), and to catalog United Greeneries (Facility #13), MedReleaf (Facility #35), Mettrum (Facility
existing cannabis strains based on metabolomic fingerprinting #41), WeedMD (Facility #60), and Canopy Growth (Facility #69). Values are
current as of April 2018.
methods and chemotaxonomy (Hazekamp et al., 2004, 2016;
Hillig, 2004; Hillig and Mahlberg, 2004; Fischedick et al., 2010;
Hazekamp and Fischedick, 2012). Some, but substantially less,
research has investigated the impact of production methods on light intensity and photoperiod (Chandra et al., 2011a, 2015),
yield and cannabinoid profiles. This includes a study on the temperature (Chandra et al., 2011b), fertilization (Malceva et al.,
use of microbial inoculants (Winston et al., 2014), the role of 2011; Caplan et al., 2017), physiological stresses (Lydon et al.,

1987; Marti et al., 2014) and elicitors (Flores-Sanchez et al., with excessive missingness (CO2 concentration (ppm), light
2009; Mansouri et al., 2009a,b, 2011, 2013, 2016; Mansouri and intensity (PAR µmol m−2 s−1 ), fertilizer rate or inoculation
Asrar, 2012). These strategies have all played an important role in with Mammoth PTM ) were not considered. Tmax , photoperiod
closing the yield gap in other crops and should be considered a during vegetative growth and duration of the vegetative or
good starting point for cannabis research. flowering periods were highly correlated to other variables (|r|
In this meta-analysis, we examine the role of plant variety > 0.75) and were therefore not included in the analysis. Prior to
(genotype) and production conditions (plant density, light, analysis, the remaining variables were standardized (mean = 0
fertilizer, temperature and duration of the flowering growth and standard deviation = 1) using PROC STANDARD and
stage) on yield per plant, per square meter and per W of lighting, categorical variables (light type, fertilizer type or variety) were
and THC and CBD yield per plant and per square meter. recoded as binary variables (0 or 1).
We describe currently available genomics and transcriptomics PROC REG, with the SELECTION = STEPWISE option,
data for cannabis and how these can be used to produce a was used to stepwise select variables. The list of unselected
better understanding of the cannabis plant. We also examine variables included the experimental continuous variables (plant
the role of production conditions in predicting plant yields and density, light intensity, duration of the flowering period, and
examine the potential use of light emitting diodes and plant pot size) and their squared effects, categorical variables (light
growth promoting rhizobacteria as novel production methods for type, fertilizer type, and variety), and the cross-products between
obtaining high yields. the continuous and categorical variables. Models were then
constructed using PROC GLIMMIX with stepwise selected
MATERIALS AND METHODS variables. A distribution to model the residuals was selected by
comparing model fit statistics between gamma, inverse Gaussian,
Data Collection shifted-t distribution, exponential, normal, and lognormal
Data were collected as treatment means, based on variety, distributions and the model with the lowest Bayesian information
plant density (plants m−2 ), concentration of CO2 during criterion was selected. A random component was added to
cultivation, light intensity (W m−2 and photosynthetically active account for the source of the data. Components of the models
radiation, PAR, µmol m−2 s−1 ), light source (high pressure that were not statistically significant (F-test p > 0.05) were
sodium, HPS, or fluorescent), photoperiod during vegetative removed sequentially until all variables remaining in the model
growth and flowering stage (h), maximum temperature during were statistically significant. In some models, numerical class
growth (◦ C), and fertilizer rate (mg N L−1 ) from Vanhove et al. variables were classified as categorical variables to produce
(2011, 2012), Potter and Duncombe (2012), Potter (2014), Caplan estimates for least squares-means.
et al. (2017) and Conant et al. (2017) (Table S2). Based on
availability, yield was recorded as either yield per plant, yield m−2
and/or yield W−1 ; percent THC and CBD in flowers at harvest RESULTS
were also recorded (Table S1). For data obtained from Potter
and Duncombe (2012), yield m−2 was calculated by multiplying Models were constructed to describe yield plant−1 , yield m−2 ,
yield W−1 (g W−1 ) by light intensity (W m−2 ); yield plant−1 yield W−1 , THC and CBD yield plant−1 and m−2 . Given the
was calculated by multiplying yield m−2 by plant density (plants high correlations, the effects of density cannot be separated
m−2 ). For data obtained from Vanhove et al. (2011, 2012), from the effects of maximum temperature during cultivation
yield W−1 was calculated by dividing yield m−2 (g m−2 ) by and the photoperiod used during the vegetative growth period.
light intensity (W m−2 ). For data obtained from Vanhove et al. Therefore, the effect of maximum temperature is interpreted as
(2011) and Potter and Duncombe (2012) THC yield (mg plant−1 ) having the same effects as plant density, whereas the vegetative
was calculated by multiplying the proportion of THC in plant photoperiod had the inverse effect as density. Likewise, the effects
material (percent divided by 100) by the yield plant−1 (mg). For of maximum temperature and duration of the vegetative growth
data obtained from Vanhove et al. (2011), THC yield m−2 (mg period have effects that are the inverse of flowering duration
m−2 ) was calculated by multiplying the proportion of THC in effect. Because yield m−2 and W−1 , THC m−2 and CBD m−2
plant material (percent divided by 100) by the yield m−2 (mg). are most relevant for industry, those results are highlighted here.
For data obtained from Vanhove et al. (2011), CBD yield (mg Formulae to predict yield, THC and CBD plant−1 are found in
plant−1 ) was calculated by multiplying the proportion of CBD in the Supplementary File.
plant material (percent divided by 100) by the yield plant−1 (mg) Based on the studied data, yield m−2 can be predicted using
and CBD yield m−2 (mg m−2 ) was calculated by multiplying the the formula:
proportion of CBD in plant material (percent divided by 100) by
the yield m−2 (mg). For yield W−1 data obtained from Potter 1 −6 −6

2 = 5.136 × 10 + −1.66 × 10 × Fdur
and Duncombe (2012), data was extracted from figures using Yield m−2
WebPlotDigitizer software (available at https://apps.automeris. 
Ltype VSS b2

io/wpd/).  1 1 5.545 × 10−6 
 
Modeling Approach  1
+  0 0.00022 

 0 1 −5.78 × 10−7 
Data used for analysis can be found in Table S3. All analyses
0 0 0
were conducted using SAS 9.4 (SAS Institute Inc. 2013). Variables

higher yields W−1 compared to other varieties at 600 W m−2 and

Early Pearly was less sensitive to this decrease compared to other
varieties (p = 0.0006 and p = 0.0099, respectively) (Figure 4A).
While increasing plant density reduced yield W−1 , the effect was
slightly different for G1 and White Widow compared to other
varieties (p = 0.0133 and p = 0.0042, respectively) (Figure 5A).
Yield W−1 was higher for plants grown using slow release
fertilizer compared to the CannaTerra nutrient regime (p < 0.05)
and when slow release fertilizer was applied, White Berry had
higher yield W−1 than other varieties (p < 0.05) (Figure 6).
For plants fertilized with CannaTerra, increased light intensity
increased yield W−1 (p < 0.0001). Yield W−1 increased with
flowering duration and this effect was stronger for the variety
Northern Lights #5 × Haze than other varieties (p = 0.0013).
THC per m−2 can be described according to:
FIGURE 2 | Effect of light type on cannabis yield per square meter. High
pressure sodium (HPS) lamps produce higher yields than metal halide (MH)   
VWa b2
lamps and Super Skunk plants produce higher yields than other varieties when

ln THC m−2 = 11.1634 + (0.1397 × Lint ) + D ×  1 −0.1108
grown under MH lamps.
0 0.2274
     
VWW b3 VBB b4
+ D ×  1 −0.3824 + Lint ×  1 0.4040
where Fdur is duration of the flowering period on the statistically 0 0 0 0
standardized scale, Ltype is light type (where 0 = HPS and  
VNLX b5
  
VWW b6

1 = MH) and VSS = 1 indicates Super Skunk. For varieties other + FD ×  1 1.2069 + Ps ×  1 −0.1676
than Super Skunk, plants grown under HPS lamps had higher 0 0.7397 0 0.1735
yields m−2 than plants grown under MH lamps (p < 0.0001)  

Ftype VSS b8

and for other varieties grown under MH lamps, yields from Super VEP Ftype b7
  0 1 0.1692 

 1 1 −0.5042
Skunk plants were higher than for all other varieties (p = 0.0058)

+  + 0 0 0 
 0 0 0   
(Figure 2). Yield m−2 increased with increasing duration of the  1 1 0.4660 
0 1 0
flowering period (p = 0.0005) (Figure 3A). 1 0 0
Yield W−1 can be predicted using the formula:
  
VG1 b1
Yield W −1 = 1.0032 + D ×  1 −0.2258 where Lint is light intensity on the statistically standardized scale,
0 0.04358 D is the plant density on the statistically standardized scale,
 
VWW b2
  
VSH9 b3
 VWa = 1 indicates Wappa, VWW = 1indicates White Widow,
+ D ×  1 −0.1799 + Lint ×  1 0.2119 VBB = 1 indicates Big Bud, FD is the duration of the flowering
0 0 0 0 period on the statistically standardized scale, VNLX = 1indicates
 
VEP b4
  
Ftype b5
 Northern Lights #5 × Haze, PS is the pot size on the statistically
+ Lint ×  1 0.2192 + Lint ×  1 0.3377 standardized scale, Ftype is fertilizer type (where 0 = CannaTerra
0 0 0 0 and 1 = slow release fertilizer) and VEP = 1 indicates Early
  

Ftype VWB b7
 Pearly. THC m−2 was lower at a light intensity of 400 W m−2
VNLX b6  0 compared to 270 or 600 W m−2 (p = 0.0001) and this effect
0 −0.2848
+  FD ×  1 0.2747  + 
was stronger for Big Bud than for other varieties (p = 0.0116).

 1 1 0.4036 
0 0.09662
1 0 0 Increasing the duration of the flowering period led to increased
THC m−2 for varieties other than Northern Lights #5 × Haze
where D is plant density on the statistically standardized scale, (p = 0.0006). Increased plant density reduced THC m−2 for all
VG1 = 1 indicates variety G1, VWW = 1 indicates White Widow, varieties; this effect was stronger for White Widow than the other
Lint is light intensity on the statistically standardized scale, varieties (p = 0.0002) (Figure 5B). Increasing the pot size from
VSH9 = 1 indicates Silver Haze #9, VEP = 1 indicates Early Pearly, 5 to 11 L reduced THC m−2 for White Widow but had a much
Ftype is fertilizer type (where 0 = CannaTerra and 1 = slow smaller effect on other varieties (p = 0.0035) (Figure 7). Early
release fertilizer), FD is duration of the flowering period on the Pearly produced lower THC m−2 compared other varieties when
statistically standardized scale, VNLX = 1 indicates Northern slow release fertilizer was applied (p = 0.0004) whereas for Super
Lights #5 × Haze and VWB = 1 indicates White Berry. Increasing Skunk produced more THC m−2 compared to other varieties
light intensity reduced yield W−1 but Silver Haze #9 produced when either fertilizer was applied (p = 0.0017).

FIGURE 3 | Effect of the duration of the flowering growth period on yield and THC per square meter. Both yield per square meter (A) and THC per square meter (B)
increased with increasing duration of the flowering period. Duration of the flowering period had a strong (|r| > 0.7) negative correlation to maximum temperature and
duration of the vegetative growth period; therefore, these predictors have the opposite effects on yield and THC per square meter as duration of the flowering period.
CBD m−2 can be described according to:

 
Lint VWW b1
 400 1 −0.9719 
 
ln CBD m−2 = 7.2498 + 

400 0 −0.8820 

600 1 −1.0370 
600 0 0
where Lint is light intensity (W m−2 ) and VWW = 1 indicates

White Widow. White Widow responded differently to light
intensity than other varieties (p = 0.0077); White Widow had
lower CBD m−2 compared to other varieties at a light intensity
of 600 W m−2 , however this effect was not statistically significant
(p > 0.05) (Figure 4B).
DISCUSSION
Effect of Production Conditions on Yield
and Cannabinoid Content
As highlighted in the data presented, yields obtained for
cannabis are highly variable depending on variety, production
conditions and production methods. Furthermore, these
data highlight the discrepancy of yields obtained in industry
compared to experimental settings. This stresses the importance
of replicating industrial growing conditions in a research
setting to allow for translation to the commercial grower
setting. This applies equally to studies designed to enhance
production based on traditional methods such as fertilization,
lighting regimes, plant density and also to novel methods
to be tested, including the use of plant-growth promoting
rhizobacteria (PGPR) or LED-based lighting systems. This
FIGURE 4 | Effect of light intensity on cannabis yield per W and CBD per section describes what is currently known about cannabis
square meter. (A) Increasing light intensity reduces yield per W and this effect cultivation in the scientific literature, with some references
is stronger for most varieties other than Early Pearly and Silver Haze #9, which to industry norms, and also underlines areas of significant
maintained higher yields at 600 Wm−2 . (B) Varieties other than White Widow
opportunity for scientific development relevant to the
produced significantly more CBD at 600 Wm−2 compared to 400 Wm−2 .
cannabis industry.

FIGURE 5 | Effect of plant density on yield per W and THC per square meter. Yield per W (A) and THC per square meter (B) declined with increasing plant density.
These effects were stronger for White Widow than for other varieties of cannabis. G1 had higher yields per W compared to other varieties at a plant density of 10.
Plant density had a strong (|r| > 0.7) positive correlation with maximum temperature during cultivation and a strong negative correlation with the duration of the
vegetative photoperiod.
square meter and per W of lighting. While increasing plant

density reduced yield per W and THC per square meter,
plant density was not an effective predictor of yield per
square meter (Figure 5). The experimental designs used
cannot quantify the relative contribution of increasing
maximum temperature and/or shortening of the vegetative
photoperiod compared to increasing plant density; these
factors should be studied in more detail in future experiments.
Furthermore, Chandra et al. (2008) recorded a maximum
rate of photosynthesis for C. sativa grown at 30◦ C, compared
to plants grown at 20–40◦ C, which explains how yield per
square meter are maintained even at higher temperatures.
Furthermore, the slightly stressful conditions of increased
plant density and maximum temperature may contribute
to increased THC accumulation. Previously, accumulated
THC increased in response to the application of abscisic acid,
FIGURE 6 | Slow release fertilizer produced higher yields per W compared to
the CannaTerra fertilizer regime. When slow release fertilizer was applied,
a plant stress hormone (Table 1) (Mansouri et al., 2009a,
White Berry produced higher yields per W compared to other varieties. Mansouri et al., 2012). Increasing pot size reduced THC
per square meter, especially for the variety White Widow
(Figure 7).
Interestingly, fertilizer type (CannaTerra compared to
slow release fertilizer) affected yield per W (Figure 6)
and THC per square meter but did not affect yield per
plant or per square meter. This result are likely due to
differences in nutrient concentration, the balance of plant
nutrients, timing of application highlighting the important
need to develop adequate nutrient regimes for cannabis.
Caplan et al. (2017) provided the first publication on
this topic and demonstrated that when a liquid organic
fertilizer (4.0N - 1.3P - 1.7K) was applied at a rate of
389 mg N L−1 and 418 mg N L−1 during the vegetative
FIGURE 7 | Increasing pot size from 5 to 11 L reduced THC per square meter growth stage, yield and THC concentration in dry flower
more for White Widow compared to other varieties.
biomass were optimized, respectively, for container-
grown “OG Kush × Grizzly” plants on two coir-based
substrates. Future studies should examine the effects of
Plant Density, Pot Size and Fertilizer individual plant nutrients and their interactions on crop
Regime Affect Yield per W and THC Yield and cannabinoid yields, and studies should be expanded to
The results of the meta-analysis highlight the impact of include a wider range of cannabis growth stages, varieties and
production conditions on cannabis yield per plant, per growing substrates.

TABLE 1 | Elicitors that have been tested on cannabis and their effects on secondary metabolite concentrations, in particular THC and CBD.
Elicitor Elicitor Main result Form of C. sativa References

concentration
Yeast extract 10 mg mL−1 Shifts in metabolites were observed but cannabinoid Hairy root cell Flores-Sanchez et al., 2009
biosynthesis appeared to be absent culture
Pythium aphanidermatum 4 and 8 g mL−1
Botrytis cinerea 4 and 8 g mL−1
Salicylic acid 0.3, 0.5, 1 mM
Methyl jasmonate 0.3 mM
Jasmonic acid 100 µM
Cannabis pectin extract 84 µg mL−1
Cannabis pectin hydrolyzed 2 mL aliquot
Pectin 0.1 mg mL−1
Sodium alginate 150 µg mL−1
AgNO3 50 and 100 µM
CoCl2 -6H2 O 50 and 100 µM
NiSO4 -6H2 O 50 and 100 µM
UV 302 nm 30 s
UV 366 nm 30 min
Absisic acid 1, 10 mg L−1 Increased THC Whole plants Mansouri et al., 2009b
1, 10 µM Increased cannabichrome, cannabinol Whole plants Mansouri and Asrar, 2012
Cycocel 500, 1000, Increased/decreased THC, CBD depending on tissue, Whole plants Mansouri and Rohani, 2014
1500 mg L−1 treatment concentration, plant sex
Ethephon 1, 5, 10, 100 µM Increased cannabinoids in male and female plants Whole plants Mansouri et al., 2013
1, 5, 10, 100 µM Increased THC, decreased CBD Whole plants Mansouri et al., 2016
Gibberellic acid 5, 10, 30, 70, Increased THC, CBD Whole plants Mansouri et al., 2011
100 µM
50, 100 µM Decreased THC Whole plants Mansouri et al., 2009a
Mevinolin 0.1, 1, 10 µM Decreased THC Whole plants Mansouri and Salari, 2014
Light Intensity, Quality, and Duration of the the growing system. Literature values report yields of 0.3122–
Flowering Period Affect Flower and 1.972 g W−1 , and are influenced by strain, light intensity and
Cannabinoid Yield plant density (Toonen et al., 2006; Vanhove et al., 2011, 2012;
Yield per square meter was higher when HPS lamps were used Potter and Duncombe, 2012). Furthermore, plants, including
than when MH lamps were used (Figure 2). This is likely due cannabis, are sensitive to the spectral composition of their
to the lower luminous efficiency (i.e., lower light output per source of light, which elicits specific effects on photosynthesis,
W) for MH lamps than HPS lamps (Eichhorn Bilodeau et al., photomorphogenesis, phototropism, and photonasty (Tamulaitis
2019). This results in lower photosynthetic photon flux density et al., 2005; Hogewoning et al., 2010). Use of electrical lighting
(PPFD) for MH than HPS lamps, even if the W m−2 of the lamp systems with different spectral outputs is common in plant
is equivalent. THC and CBD per square meter increased with research and greenhouse horticulture. Most commonly, high
light intensity while yield per W decreased with increasing light pressure sodium (HPS) gas discharge lamps and fluorescent
intensity (Figure 4). The increased accumulation of THC and tubes are used (Hogewoning et al., 2010). Although the
CBD at 600 W m−2 suggest that these compounds are produced spectral emissions of these lights span the entire spectrum of
to limit the effects of light stress at higher light intensities as sunlight, they feature distinct wavelength patterns (Hogewoning
a result of a stress response (Mansouri et al., 2009b; Mansouri et al., 2010). HPS lights generally emit light most strongly
et al., 2012). Our results also clearly indicate that increasing the in the yellow-red end of the spectrum, which is absorbed by
duration of the flowering period (or reducing the duration of the chlorophyll and used in photosynthesis. Improvements in the
vegetative period) increases yield per square meter and THC per blue component of HPS lights can improve light suitability for
square meter (Figure 3), a result which Vanhove et al. (2012) plant growth, however modifications are required to optimize
attributed to increased photosynthetic assimilation directed to the red spectrum of their emissions to enhance plant growth
bud growth instead of stem and leaf growth. (Tamulaitis et al., 2005). These changes would reduce the energy
Light quality, intensity, source and photoperiod play a lost as infrared radiation or heat. In contrast, fluorescent tubes
critical role in yield and quality of cannabis. Often, yield have peaks throughout the spectrum but lack emissions in
is reported as g W−1 , as a measure of energy efficiency of the far-red region of the spectrum (Tamulaitis et al., 2005).

High power light emitting diodes (LEDs) are an emerging The effects of LEDs on plant growth and photomorphogenesis
versatile electrical light source offering many advantages over has been studied in plant species other than cannabis, with
conventional electrical light sources, including high energy emphasis on the control of flowering and/or the duration of
efficiency, long life, and especially, the possibility to test the blooming period. Physiological studies have shown that light
the effects of many spectral combinations of wavelengths on quality, quantity and duration regulate flowering (Bula et al.,
plant growth and development. This could eventually lead to 1991; Tennessen et al., 1994). According to Guo et al. (1998) and
determination of the ideal light emission spectrum, allowing Thomas and Vince-Prue (1996), red light can inhibit flowering
for lighting system designs tailored to enhance plant growth via red-light receptors such as phytochromes, which absorb light
while minimizing associated energy costs (Tamulaitis et al., 2005). effectively at wavelengths above 600 nm (Kelly and Lagarias,
In the meantime, studies have begun to exploit the spectral 1985). In contrast, blue light can inhibit flowering via blue-
elasticity of LEDs to examine the effects of different wavelength- light receptors such as cryptochromes, which absorb light well at
light combinations on plant growth. The possibility of achieving wavelengths below 500 nm (Lin et al., 1995; Banerjee et al., 2007;
higher irradiance at isolated wavelengths of light than with Eichhorn Bilodeau et al., 2019).
monochromatic light previously obtained through filters, could A study on cannabis demonstrates that flowering time is
allow more accurate assessments of plant physiological responses determined by photoperiod: flowering is induced when day
(Lefsrud et al., 2008). length is shorter than 12 h (Potter, 2014). While light quality
The optimal spectrum of light to achieve optimal yields influences on cannabis flowering have not yet been studied, light
of cannabis and cannabinoids remains to be fully elucidated. quality has been shown to influence flowering and duration of
Environmental factors, such as temperature and irradiance levels, the blooming period in marigold (Tagetes erecta L. cv. Orange
can have strong effects on plant growth and the accumulation Boy) and salvia plants (Salvia splendens F. Sello ex Ruem &
of pigments critical for photosynthesis (Lefsrud et al., 2005, Schult. cv. Red Vista) (Heo et al., 2002). The number of visible
2006). Chandra et al. (2008) discussed photosynthetic and flower buds in marigold was approximately five times higher
water-use efficiency responses of cannabis to light, CO2 and in the presence of fluorescent light (with or without red LED)
temperature levels. The study demonstrated that maximum rate than under monochromic blue or red light. Monochromic blue
of photosynthesis occurs at 30◦ C, 750 µmol CO2 mol−1 , and or red light were found to suppress bud formation in salvia
under 1,500 µmol m−2 s−1 . The study concluded that high while fluorescent light plus far-red light was also found to inhibit
intensity lighting, in drier and CO2 enriched environments flower bud formation in marigold. Day-extension using red or
promotes higher photosynthetic activity, water use efficiency, blue LEDs inhibited flower and bud appearances. Night-break
and nearly constant internal to ambient CO2 concentration treatment with red LEDs also delayed flower bud appearance in
in cannabis. okra (Abelmoschus esculentus L. Moench) and a cultivar of native
Another challenge associated with lighting systems is that rosella (Abelmoschus moschatus ssp. tuberosus Span Borss). Night
light intensity decreases with depth within the plant canopy as break with green light delayed flowering more strongly than blue
leaves absorb the light (Massa et al., 2005). In HPS and overhead light, but slightly less than red light (Hamamoto and Yamazaki,
LED lighting systems, the top of the canopy is often light 2009). In long-day plants, experiments suggest that flowering
saturated, yet the canopy as a whole is light limited. Providing is promoted most when red light is delivered during the early
additional light to the lower canopy increases the proportion part of the photoperiod and far red light toward the end of the
of light used for photosynthesis without exceeding the point of photoperiod (Lane et al., 1965; Evans, 1976; Kadman-Zahavi and
photosynthetic light saturation (Massa et al., 2005). Unlike HPS Ephrat, 1976; Thomas and Vince-Prue, 1996). However, cannabis
lamps, LEDs emit little heat and can be placed close to the crop is a short-day plant, so it remains unclear whether these results
without burning leaves, meaning they are a practical interlighting are relevant for cannabis production.
system in commercial settings. For example, LEDs located within
a cowpea (Vigna unguicultata L. Walp.) canopy improved
biomass production by 33 %, compared to plants grown under Effect of Variety on Crop Yield and
overhead lights; intercanopy lights were also associated with an Cannabinoid Content
increased energy conversion rate (Massa et al., 2005). Hawley The results of the meta-analysis show that yield per square
(2018) demonstrated that supplemental sub-canopy lighting meter and per W and accumulation of THC and CBD vary
(SCL) can increase cannabis bud yield and modify cannabinoid based on plant variety. Sawler et al. (2015) showed that variety
and terpene profiles. The increase in bud yield is assumed to name does not always correspond to genotype, as so it is
be related to increased photosynthetic photon flux densities critical that future reports, document the genotype used to
(PPFD) compared to production with overhead lighting alone. allow for comparison of results from different studies. It is
Red and blue SCL yielded a more consistent metabolite profile also worth highlighting that while Silver Haze #9 stands out
throughout the canopy, whereas red, green and blue SCL as a top-yielding variety, it was pruned differently than other
had the greatest impact on metabolite upregulation. A light varieties included in the same study. Therefore, the high yields
spectrum with comparatively more green light drove plants of this variety may be related to pruning rather than to its
to produce more carotenoids to manage green wavelengths, genotype and both possibilities should be investigated in future
and consequently up-regulated other related terpenes research. Our results confirm the findings of Vanhove et al.
in the process. (2012), who showed that varieties respond differently to changes

in production conditions, as evidenced by multiple significant been deposited in NCBI, including whole genome sequences,
variety-by-production condition interaction effects. genotype by sequence, and short read assemblies. Many of
these accessions are not associated with publications, and lack
Cannabis Genetic and Chemical Diversity metadata to permit their full use by the research community.
Cannabis plants are be classified as indica, sativa, and In spite of the lack of metadata, these genome accessions can
ruderalis. Lack of scientific consensus means these terms be used to examine variation in the genomes of a range of
refer to cannabinoid content, morphology, allele frequencies cannabis cultivars.
or provenance (Hillig, 2005; Dufresnes et al., 2017). The first published cannabis transcriptomes were synthesized
Historically, hemp-type (high in cannabidiolic acid, CBDA) from the roots, stems, vegetative shoots, pre-flowers and flowers
and medical/recreational-type (often called marijuana, high of Purple Kush; more than 18.8 Gb of poly-A+ RNA reads
in tetrahydrocannabinolic acid, THCA) strains have been corresponding to 30,000 genes were identified (Van Bakel et al.,
categorized by their chemotype. For example, hemp is legally 2011). Since then, a leaf tissue salinity response transcriptome has
defined by EU and Canadian regulations as containing <0.3% also been published (Liu et al., 2016). A slightly larger number of
THC (Canada, 1998). Species level classification of Cannabis transcriptome studies exist for hemp-type cannabis plants (Behr
plants is complicated by the lack of reproductive barriers between et al., 2016; Booth et al., 2017; Guerriero et al., 2017). However,
individuals conventionally described as subspecies, phenotypic the functional characterization of the cannabis genome is still in
plasticity, strong artificial selection for fiber-type and drug-type its infancy.
plants, as well as mixing of wild and cultivated populations
since antiquity (Sawler et al., 2015; Clarke and Merlin, 2016; Crop Improvement Using Genomics
Grassa et al., 2018). More recently, genomic and transcriptomic and Transcriptomics
distinctions between hemp and medical/recreational cannabis The diverse uses of cannabis plants are reflected in the
have been made (Piluzza et al., 2013; Sawler et al., 2015). Sawler significant variation in their stalk height, seed size, fiber length,
et al. (2015) identified ∼14,000 single-nucleotide polymorphisms phytochemical concentrations, and sensitivity to day length
that distinguished hemp-type and medical/recreational-type (Clarke and Merlin, 2016). Many of these traits, including
plants. Welling et al. (2016a) used genomic markers to predict those typically attributed to indica, sativa and ruderalis-
the cannabinoid profile of 22 Cannabis accessions with over type plants (Gould, 2015), may be targeted for improvement
98% accuracy, thereby confirming the genetic underpinning using conventional or modern breeding technologies. Detailed
of chemotype. knowledge of the variation that exists across the Cannabis genus
The Cannabis genome is diploid (2n = 20) with nine is fundamentally important to any project aiming to improve
autosomal chromosome pairs and one pair of XY sex cultivars. Several projects have characterized the genetic structure
chromosomes (Sakamoto et al., 1998; Divashuk et al., 2014). of small populations of cannabis (Gao et al., 2014; Sawler
The nuclear genome was characterized and determined to be et al., 2015; Soorni et al., 2017), but this has not yet been
∼1,636 Mb for female plants (XX) and 1,683 Mb for male plants done on a larger scale. This synthesis of the knowledge has
(XY) (Sakamoto et al., 1998). In 2011, the first draft haploid not yet transpired as the illicit nature of the drug-type plant
genome sequences were published (Van Bakel et al., 2011). These has delayed the establishment of a well-conserved and well-
included a female clone of the drug-type cultivar Purple Kush, annotated germplasm with consistent nomenclature (Clarke and
and a female plant of the fiber-type cultivar Finola (Van Bakel Merlin, 2016). There is a movement in the cannabis research
et al., 2011). Theses genomes were assembled from Illumina community to preserve and analyze germplasm across the genus
paired-end (6 libraries with median insert sizes ranging from to facilitate research and breeding programs (Clarke and Merlin,
220 to 600 bp), Illumina mate-pair (2 libraries with median 2016; Welling et al., 2016b; Small, 2018).
insert sizes of 1.8 and 4.6 kb), and 454 mate-pair (11 libraries Starting in the 1990s, molecular markers for cannabis varieties
with median insert sizes ranging from 8 to 80 kb) libraries (Van were developed for forensic analysis of plant origin. Hemp
Bakel et al., 2011). The assembled Purple Kush genome was breeders have since integrated molecular markers (namely sex-
786.6 Mb including 252 Mb of gaps. The presence of gaps in linked and chemotypic markers) to enhance marker-assisted
the genome was attributed to high repeat content and to high selection (MAS) strategies for crop improvement (Mandolino
sequence variation in the cannabis genomes. More recently, an and Carboni, 2004; Faux et al., 2016) and cannabis researchers
ultra-high-density genetic map was generated for Cannabis using have used QTL analysis to identify loci associated with
a combination of long and short read sequencing technologies THCA production (Weiblen et al., 2015). A number of
across parental, F1, and 96 recombinant F2 individuals (Grassa marker sets have been generated for a variety of genetic loci
et al., 2018). Long-read technologies, including those from including microsatellites (Dufresnes et al., 2017) and SNPs
PacBio, have been used to sequence through repetitive regions, associated with traits of interest (Sawler et al., 2015; Lynch
in order to close sequencing gaps in a number of plant species. et al., 2016; Soorni et al., 2017). Following the advent of
Several long-read cannabis genome sequences have been next-generation sequencing, QTL mapping and genome-wide
contributed to the NCBI Genome repository. None of these association studies have become more feasible, which will
sequences are associated with peer-review publications, nor are accelerate the discovery of important markers. Due to the high
they presented as assembled or annotated genomes. Additionally, phenotypic plasticity of cannabis, associations between markers
more than 1,500 short-read genome sequencing samples have and phenotypes must be carefully characterized (Salentijn et al.,

2015). The advent of genome editing technologies also hold great greenhouse production. However, producers are beginning to
promise for cannabis improvement as Agrobacterium mediated produce cannabis for the recreational market under greenhouse
transformation protocols have been published (Feeney and conditions, as it allows for larger cultivation areas and the
Punja, 2003). use of natural sunlight, which reduces heating and lighting
Efficient genome editing capabilities facilitated by the costs. To date, literature is scarce around best practices for
biotechnologies of CRISPR-Cas9 and related technologies cannabis growing methods. Several cultivation methods are
hold great promise for targeted improvement of Cannabis used within growing rooms, including traditional bench setups,
cultivars. For these technologies to be implemented aeroponics, and hydroponics. While, Potter (2014) reviewed
three companion methodologies must be established: (1) growing conditions used in industry they did not provide
micropropagation; (2) efficient transformation; (3) plant comparisons of productivity based on growing methods. While
regeneration. Micropropagation technologies are foundational growers are keen to obtain high yields in each growth cycle,
to the Cannabis industry, where they are used primarily with the another challenge is the ability to obtain the maximum number
aim of propagating and expanding high value cannabis plants. of growing cycles per year (personal communication).
For the purposes of biotechnology applications, it is necessary With the adoption of the Cannabis Act, Health Canada
to develop and maintain cultures pluripoten stem cells as callus regulations will allow for outdoor cultivation of cannabis. While
or cell suspension culture. Since the 1970s, a number of such differences certainly will exist, producers interested in outdoor
protocols have been established for Cannabis (reviewed in Lata production of cannabis could adopt knowledge developed for
et al., 2017; Wróbel et al., 2018). Transformation of Cannabis cells agronomic practices (fertilization, seeding rate, harvest time,
using Agrobacterium tumafasciens and A. rhizogenes have been etc.) for cultivation of hemp (Atal, 1961; Mechtler et al.,
demonstrated starting in 2003 with the transformation of callus 2004; Amaducci et al., 2008; Cosentino et al., 2012; Faux
(Feeney and Punja, 2003) and more recently using callus derived et al., 2013; Finnan and Burke, 2013a,b; Faux and Bertin,
from a variety of tissue types and cultivars (Slusarkiewicz-Jarzina 2014; Aubin et al., 2015, 2016; Razumova et al., 2016).
et al., 2005; Wahby et al., 2013). Protocols for the transformation However, it remains unclear how these conditions will influence
of Cannabis roots have also been established (Wahby et al., 2013). medical/recreational cannabis quality aspects (flower yield,
The primary and persistent challenge has been to regenerate cannabinoid concentration), which are different from hemp
plants from the transformed callus and explant tissue (Feeney quality variables (seed yield, fiber content). Thus, these factors
and Punja, 2003); plant recovery rates range from <2% to more will need to be investigated in the context of field cultivation
than 50% (Chaohua et al., 2016) depending on the protocol, of medical/recreational cannabis. The remainder of this section
starting tissue, and genotype used. To date, we are not aware of focuses on factors that affect cannabis yield and quality in the
any published accounts of CRISPR mediated genome editing in context of indoor, controlled environment production.
Cannabis; this will undoubtedly not be the case for long.
Limitations of Available Cannabis Data Potential Role for Plant-Growth Promoting

This meta-analysis was able to identify some key factors that Rhizobacteria in Cannabis Production
contribute to cannabis yields. However, only three studies were The role of the phytomicrobiome in regulating plant growth
included in the meta-analysis due to the fact that other published has received significant attention in the recent scientific
studies did not report sufficient information about growing literature and has been the basis for many crop-yield-
conditions for inclusion in the models (Table S2). Furthermore, enhancing technologies (e.g., Backer et al., 2018). Several
it remains difficult to determine the relationship between flower studies have surveyed the diversity of bacterial and fungal
and cannabinoid yields due to the lack of consistent reporting endophytes in medical/recreational cannabis and hemp
of cannabinoid concentration or yield. Our results also show and have found that colonization depends on the cannabis
that light type, as a proxy for PPFD, has a significant impact genotype, the plant tissue sampled and the timing of sample
on flower and CBD yields, which suggests that reporting of light collection relative to the plant growth stage. Among plants
intensity as W m−2 is insufficient on its own. Finally, the results sampled from India, Pakistan, the USA and Canada the most
of this meta-analysis show that yield per square meter obtained common bacterial genera associated with medical/recreational
in scientific studies (Table S1) remains much lower than yield cannabis and hemp plants were Pseudomonas, Staphylococcus,
per square meter obtained in industry (Figure 1) suggesting that Bacillus, Acinetobacter, Chryseobacterium, Enterobacter, and
discrepancies remain between industry production practices and Microbacterium while Erwinia, Cedecia, Chryseobacterium,
growing conditions used in scientific studies. This highlights the Enterobacter, Microbacterium were found but at lower
value of knowledge exchange between academia and industry. frequencies (Gautam et al., 2013; Winston et al., 2014; Afzal
et al., 2015; Scott et al., 2018). These studies also determined
Future Considerations in that the colonization frequency was highest for leaves, followed
Cannabis Research by stems and petioles, however, these studies did not consider
Diversification of Cultivation Systems for Cannabis bacteria residing in or near root tissue. Community composition
Currently, cultivation of medical cannabis is usually conducted was determined mainly by soil type while community structure
in controlled environment growing rooms since they offer a was determined by cultivar. These results highlight the need for
higher degree of control over growth conditions, compared to systemic studies of microbial diversity in cannabis, with time

points spanning from seed germination through to maturity, plants demonstrated their potential antagonistic activity against
including leaf, stem, petiole, flower, and root tissue. Aspergillus flavus, Botrytis cinereal, Ceratocystis fimbriata,
Many of the isolates identified in the studies mentioned Colletotrichum gloeosporioides, Curvularia lunata, Fusarium
above tested positively in vitro for properties associated with oxysporum, Geotrichum candidum, Fusarium solani, Rhizoctonia
plant growth promotion (siderophore, cellulose, organic acid, solani, Sclerotinia sclerotiorum, and Trichothecium roseum,
and/or indole-3-acetic acid production and/or P-solubilization). in vitro (Gautam et al., 2013; Kusari et al., 2013; Qadri et al.,
In planta, two isolates were able to increase canola (Brassica 2013; Scott et al., 2018). The second option, inducing ISR, is
napus) root length under salt stress conditions Afzal et al. of particular interest in cannabis production given (1) the high
(2015), while other isolates did not increase growth variables susceptibility of flowers to infection by plant pathogens and
of tomato (Solanum lycopersicum L.) or hemp seedlings (Scott (2) the necessity to maintain extremely low pesticide residue
et al., 2018). Bioprospecting from wild cannabis may reveal levels on flowers. However, this remains to be tested in cannabis.
PGPR that improve cannabis growth (Kusari et al., 2017). Finally, harnessing the biocontrol aspect of PGPR to clean
Alternatively, PGPR isolated from other crops may provide growing rooms in between growth cycles could reduce the risk
significant potential for improving cannabis yields. For example, of contamination between batches and reduce time between
Conant et al. (2017) reported that Mammoth PTM , a consortium growth cycles.
of P-mobilizing microorganisms, increased flower yield per plant Several studies have already investigated the role of
by 16.3% from 15.9 g (control) to 18.5 g (with Mammoth PTM ). endophytes in cannabis growth and development; while
Since P-solubilization is only one of a set of mechanisms that data are still lacking about effects on growth and yield of
microbes can use to promote plant growth, these results represent cannabis and the accumulation of cannabinoids in response to
a promising starting point and suggest that testing microbes plant inoculation with PGPR. In contrast, multiple reports have
that increase plant yield by other mechanisms is warranted. investigated the role of cannabis endophytes for biocontrol; these
Additionally, PGPR from the genus Bacillus have been shown have demonstrated strong potential for control of fungal and
to accelerate time to flowering for crops such as banana (Musa bacterial pathogens in vitro. The role of cannabis endophytes for
acuminata cv. “Berangan”), marigold (Tagetes erecta L.) and biocontrol remains to be tested in planta.
carnation (Dianthus carophyllus L.) (Mia et al., 2005; Flores
et al., 2007; Kumar et al., 2014). Achieving flowering in a shorter
timespan would reduce the time to harvest for each growth cycle
CONCLUSIONS
to help growers attain a higher number of harvests per year. In order to increase cannabis yield per square meter and per
In addition to increasing dry flower yield, inoculation with W light, the results of this meta-analysis point to the use of
PGPR has the potential to increase cannabinoid yield via (1) low plant density (≤12 plants per square meter), (2) a
elicitation; this has been previously demonstrated for secondary flowering period duration of 9 weeks, (3) the use of HPS lamps,
metabolites in other plant species (previously reviewed by (4) an adequate fertilizer regime, and (5) manipulating light
Gorelick and Bernstein, 2017). Several studies, cataloged in intensity to preserve high energy efficiency vs. favor THC and
Table 1, have tested the role of biotic and abiotic elicitors on the CBD accumulation. Furthermore, our results demonstrate that
effects of cannabinoid biosynthesis, revealing the sensitivity of cannabis varieties respond differently to production conditions.
this pathway to external signals. In contrast, studies conducted The vast amount of existing genomic and transcriptomic data
by Mansouri et al. (2009a,b, 2011, 2013 and 2016), Mansouri and can be used to catalog current cannabis diversity resulting
Asrar (2012), Mansouri and Rohani (2014) and Mansouri and from thousands of years of breeding and used to identify
Salari (2014) demonstrated that abscisic acid, cycocel, ethephon, area for crop improvement. While these basic production
gibberellic acid, and mevinolin can all alter cannabinoid conditions are further investigated, we also propose the use
biosynthesis. While Flores-Sanchez et al. (2009) tested a large of additional technologies such as LEDs to increase power-use
number of elicitors for effects on cannabis hairy root cell efficiency, and PGPR to increase nutrient efficiency and regulate
cultures, this did not induce cannabinoid biosynthesis. Testing cannabinoid yield.
the same elicitors in whole plants could lead to up- or down-
regulation of cannabinoid biosynthesis. Furthermore, it has been
demonstrated that bacteria isolated from one crop or plant AUTHOR CONTRIBUTIONS
species can stimulate growth and induce systemic resistance
other crop species (Smith et al., 2015; Fan, 2017). Therefore, RB, OW, and DS conceptualized the review layout. RB wrote
bacteria isolated from other crop species could be tested for and edited the review. PR, VM, SE, DL, and MA contributed
effects on cannabinoid biosynthesis in cannabis plants. significant portions of the text. TS conducted the meta-analysis
PGPR also offer the potential to close the yield gap by with input from RB. RB and TS produced figures and tables. GR,
reducing yield losses due to plant pathogens. PGPR can ML, OW, and DS critically reviewed the manuscript.
reduce yield losses by (1) inhibiting pathogen growth in planta
or in soil via antagonism, (2) inducing systemic resistance FUNDING
in the plant, (3) reducing contamination between growth
cycles. Strong evidence exists in the scientific literature to Funding for this work was provided by a Collaborative Research
support the first mechanism. Endophytes isolated cannabis and Development grant, Enhanced yield and cannabinoid

production of homogenous medical marijuana plants (award Greeneries, MedReleaf, Mettrum, WeedMD and Canopy Growth. Facility numbers
number 517552-17), from the Natural Science and Engineering corresponds to those shown in Figure 1 of the text. Data were collected from
press releases provided by the companies, with links provided
Council of Canada in collaboration with Ravenquest Biomed.
in column G.
ACKNOWLEDGMENTS Table S2 | Yield data for cannabis (drug type) from the scientific literature based
on growing conditions reported by authors. Units are as indicated in each column
title. HPS, high pressure sodium; MH, metal halide; org, organic fertilizer (no
The authors wish to acknowledge the editor and reviewers for further details provided by author); slowrel, slow release fertilizer contained in
their constructive feedback of this review. growing medium; Cap2017, Caplan et al., 2017; Con2017, Conant et al., 2017;
Pot2012, Potter and Duncombe, 2012; Pot2014, Potter, 2014; Van2011,
SUPPLEMENTARY MATERIAL Vanhove et al., 2011; Van2012, Vanhove et al., 2012.
Table S3 | Cannabis yield data included in the statistical analysis. Data for Caplan
The Supplementary Material for this article can be found et al. (2017), Conant et al. (2017), and Potter and Duncombe (2012) were
online at: https://www.frontiersin.org/articles/10.3389/fpls.2019. excluded from the meta-analysis due to a high degree of missing information
00495/full#supplementary-material about growing conditions. Units are as indicated in each column title. HPS, high
pressure sodium; MH, metal halide; slowrel, slow-release fertilizer contained in
Table S1 | Projected and actual yield data for commercial cannabis operations in growing medium; Pot2012, Potter and Duncombe, 2012; Van2011, Vanhove
Canada as of April 2018. Actual yields are reported for AB Labs, United et al., 2011; Van2012, Vanhove et al., 2012.
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published: 24 April 2019

Closing the Yield Gap for Cannabis: A

Frontiers in Plant Science | www.frontiersin.org 1 April 2019 | Volume 10 | Article 495

Regulations (ACMPR) and its predecessor acts: Marihuana for

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higher yields W−1 compared to other varieties at 600 W m−2 and

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CBD m−2 can be described according to:

where Lint is light intensity (W m−2 ) and VWW = 1 indicates

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square meter and per W of lighting. While increasing plant

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Elicitor Elicitor Main result Form of C. sativa References

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Frontiers in Plant Science | www.frontiersin.org 8 April 2019 | Volume 10 | Article 495

Frontiers in Plant Science | www.frontiersin.org 9 April 2019 | Volume 10 | Article 495

Limitations of Available Cannabis Data Potential Role for Plant-Growth Promoting

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Frontiers in Plant Science | www.frontiersin.org 11 April 2019 | Volume 10 | Article 495

Frontiers in Plant Science | www.frontiersin.org 12 April 2019 | Volume 10 | Article 495

Frontiers in Plant Science | www.frontiersin.org 13 April 2019 | Volume 10 | Article 495

Frontiers in Plant Science | www.frontiersin.org 14 April 2019 | Volume 10 | Article 495

Frontiers in Plant Science | www.frontiersin.org 15 April 2019 | Volume 10 | Article 495

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