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Light-emitting-diode (LED) lighting for greenhouse tomato production Paul Deram Under the supervision of Dr. Mark Lefsrud Department of Bioresource Engineering Macdonald Campus McGill University, Montréal January, 2013 A thesis submitted to McGill University in partial fulfillment of the requirements of the degree of Master of Science. ©Paul Deram 2013
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Page 1: Light-emitting-diode (LED) lighting for greenhouse tomato production

Light-emitting-diode (LED) lighting for greenhouse tomato

production

Paul Deram

Under the supervision of Dr. Mark Lefsrud

Department of Bioresource Engineering

Macdonald Campus

McGill University, Montréal

January, 2013

A thesis submitted to McGill University in partial fulfillment of the

requirements of the degree of Master of Science.

©Paul Deram 2013

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ABSTRACT

The cost of artificial lighting is a major expense in the greenhouse production

industry, especially during the winter where supplemental lighting is required to

maintain production. Current technology uses broad spectrum high pressure

sodium lamps (HPS), which, despite being excellent luminous sources, are not the

most efficient light source for plant production. Specific light frequencies have

been shown to impact photosynthesis more directly than others (especially in the

red and blue ranges); focusing on specific wavelengths, light-emitting diodes

(LEDs) could diminish lighting costs due to their high efficiency and lower

operating temperatures. LEDs can be selected to target the wavelengths absorbed

by plants, enabling the growers to customize the wavelengths of light required to

maximize production and limit wavelengths that do not significantly impact plant

growth. The primary purpose of this experiment was to test tomato plants

(Solanum lycopersicum), in a research greenhouse using a full factorial design

with three light intensities (High: 135 µmol m-2 s-1, Medium: 115 µmol m-2 s-1 and

Low: 100 µmol m-2 s-1) at three red to blue ratio levels (5:1, 10:1 and 19:1)

compared to 100% HPS, and a control (no supplemental lighting). The exact

wavelengths chosen were 449 nm for the blue and 661 nm for the red. Secondary

treatments were also tested using 100% red light supplied from the top, 100% red

light supplied from the bottom, a 50%:50% LED:HPS and a replicate of the 10:1

ratio with High light intensity. The experiment was replicated over two different

seasons (Summer-Fall 2011 and Winter-Spring 2011-2012). During the

experiment, the highest biomass production (excluding fruit) occurred with the

19:1 ratio (red to blue), with increasing intensity resulting in more growth,

whereas a higher fruit production was obtained using the 5:1 ratio. The highest

marketable fruit production (fruit over 90 g, Savoura internal standard) was the

50%:50% LED:HPS, followed by 5:1 High and 19:1 High. From this research,

LEDs have been shown to be superior in fruit production over HPS alone, and

LEDs can improve tomato fruit production with HPS and have the ability to

become the dominant supplemental greenhouse lighting system.

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RÉSUMÉ Le coût de l'éclairage artificiel est une dépense importante dans le secteur de la

production en serre, surtout en hiver lorsqu’un éclairage supplémentaire est

nécessaire pour maintenir le niveau de production. La technologie actuelle utilise

des lampes à haute pression de sodium (HPS), qui en dépit d'être d'excellentes

sources lumineuses, ne sont pas les sources lumineuses les plus efficaces pour la

production végétale. Certaines fréquences spécifiques de lumière ont montré avoir

un impact plus direct sur la photosynthèse que d'autres (en particulier dans les

gammes de rouge et de bleu); en mettant l'accent sur certaines longueurs d’onde,

les diodes électroluminescentes (LED) pourraient diminuer les coûts d'éclairage,

en raison du rendement élevé et des températures plus basses de ce type de lampe.

Les LED peuvent cibler les principales fréquences de lumière mieux absorbées

par les plantes, ce qui permettrait aux producteurs de créer une lumière aux

longueurs d'onde adaptées à la production optimale des plantes. Le principal

objectif de cette expérience était de tester les lampes sur des plants de tomate

(Solanum lycopersicum) dans une serre de recherche en utilisant un plan factoriel

complet avec trois intensités lumineuses (Haute: 135 μmol m-2 s-1, Moyenne: 115

μmol m-2 s-1 et Basse: 100 μmol m-2 s-1) et trois proportions de rouge et bleu (5:1,

10:1 et19: 1), et comparer leur performance à celle de 100% HPS, et d’un contrôle

(pas d'éclairage supplémentaire). Les longueurs d'onde choisies sont 449 nm

(bleu) et 661 nm (rouge). Certains traitements secondaires ont également été

testés, dont 100% rouge (éclairage par le haut ou le bas), un 50%:50% LED:HPS

et une reproduction du 10:1 à haute intensité. L'expérience a été menée au cours

de deux saisons différentes (été-automne et hiver-printemps). La production

végétative la plus importante s'est produite avec le rapport 19:1 (rouge : bleu). La

production de fruits était la plus élevé avec le rapport 5:1. La production en fruits

commercialisables la plus importante (fruits de 90 g et plus : étalon interne de

Savoura) a été pour le 50%:50% LED:HPS, suivi du 5:1 et 19:1 à haute intensité.

Les LED se sont montrés supérieures aux HPS quant à la production de tomates.

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ACKNOWLEDGEMENTS This thesis is the result of the input of numerous parties. First of all, I would like

to thank the company, General Electric Lighting Solutions Canada (Nabil Jacques

Salem, Jean-François Richard) for proposing the research project, and committing

to the research with funding and development of the LED array prototypes.

I would also like to thank my supervisor: Dr. Mark Lefsrud for providing me

with much needed support throughout the duration of this research project, as well

as opening up his laboratory for me to conduct my research, and for sharing his

personal expertise in the domain of LED lighting for plant research.

I would like to thank Dr. Valérie Orsat, for her always kind and encouraging

advice, as well as for all the help given as part of my advisory committee for the

sake of this project.

To Savoura in Portneuf, QC (David Brault and Claire Boivin), I extend my thanks

for their knowledge in tomato growing and care techniques which was

instrumental in the completion of this research, as well as for teaching me how to

work efficiently and in a timely fashion in a greenhouse operation.

To any others who helped me during the greenhouse experiments, for answering

my questions, for advice on methodology or even for their friendship, thank you

all (Alejandro Jaul, Nicholas Matlashewski, Allison Busgang, Michael Schwalb,

Julie Gagné, Anil Patel, Yvan Gariépy and many others).

Finally, I would like to thank my family, friends and my wonderful fiancée, for all

their moral support, and for making their unseen presence known and felt.

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TABLE OF CONTENTS

ABSTRACT ............................................................................................................. 2

RÉSUMÉ ................................................................................................................. 3

ACKNOWLEDGEMENTS ..................................................................................... 4

TABLE OF CONTENTS ......................................................................................... 5

LIST OF TABLES ................................................................................................... 8

LIST OF FIGURES ................................................................................................. 9

LIST OF EQUATIONS ......................................................................................... 11

ABBREVIATIONS ............................................................................................... 12

1. INTRODUCTION .......................................................................................... 13

2. HYPOTHESIS AND OBJECTIVE ................................................................ 15

3. LITERATURE REVIEW ............................................................................... 16

3.1. Tomato and Greenhouses ............................................................................ 16

3.2. Photosynthesis ............................................................................................. 17

3.3. Lighting Systems: ........................................................................................ 20

3.4. LEDs in Plant Research:.............................................................................. 23

3.5. Effects of Different Wavelengths ................................................................ 24

3.6. Effect of Intensity ........................................................................................ 27

4. MATERIALS AND METHODS ................................................................... 29

4.1. Plant Care .................................................................................................... 29

4.2. Experimental Setup ..................................................................................... 31

4.2.1. Greenhouse Setup ................................................................................. 31

4.2.1. LED Setup ............................................................................................ 34

4.3. Instrumentation ............................................................................................ 36

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4.4. Statistical Analysis ...................................................................................... 39

5. RESULTS ....................................................................................................... 41

5.1. Fruit ............................................................................................................. 41

5.1.1. Marketable Fruit Number ..................................................................... 41

5.1.2. Marketable Fruit Mass .......................................................................... 42

5.1.3. Total Fruit Number ............................................................................... 44

5.1.4. Total Fruit Mass .................................................................................... 45

5.1.5. Average Fruit Mass............................................................................... 47

5.2. Vegetative Biomass ..................................................................................... 49

5.2.1. Fresh Mass ............................................................................................ 49

5.2.2. Dry Mass ............................................................................................... 50

5.2.3. Dry to Fresh Biomass Ratio .................................................................. 52

5.2.4. Fruit and Flower Counts ....................................................................... 53

5.3. Fruit to Biomass Ratio ................................................................................. 55

5.4. Environmental Data ..................................................................................... 57

5.4.1. Temperature .......................................................................................... 57

5.4.2. Relative Humidity ................................................................................. 57

5.4.3. Light Sensor Map ................................................................................. 58

6. DISCUSSION ................................................................................................. 60

6.1. Ratios chosen ............................................................................................... 60

6.2. Fruit ............................................................................................................. 61

6.2.1. Marketable Fruit ................................................................................... 61

6.2.2. Total Fruit ............................................................................................. 63

6.3. Vegetative Biomass ..................................................................................... 64

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6.4. HPS to LED comparison ............................................................................. 66

6.5. Top versus Bottom Lighting Systems ......................................................... 66

6.6. Light Measurement Techniques .................................................................. 67

6.7. Production Issues ......................................................................................... 71

6.7.1. First Run: .............................................................................................. 72

6.7.2. Second Run: .......................................................................................... 74

6.8. Observations for future research ................................................................. 76

7. CONCLUSIONS ............................................................................................ 81

8. REFERENCES ............................................................................................... 83

APPENDIX A: Formatted Data ............................................................................. 89

APPENDIX B: Statistical Data (Box Plots) .......................................................... 94

APPENDIX C: Raw Data .................................................................................... 101

APPENDIX D: Weather and Lighting Data ........................................................ 110

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LIST OF TABLES

Table 1: Greenhouse section placement ............................................................... 31

Table 2: Treatment list and description.................................................................. 35

Table 3: End of first run and start of second run light map data............................ 38

Table 4: Fruit and flower count summary .............................................................. 54

Table C1: Total fruit data ..................................................................................... 101

Table C2: Red fruit data ....................................................................................... 103

Table C3: Marketable fruit data (> 90 grams) ..................................................... 104

Table C4: Biomass data ....................................................................................... 105

Table C5: Fruit and flower count data ................................................................. 108

Table D1: Temperature data during the second run ............................................. 110

Table D2: Temperature data during the first run ................................................. 111

Table D3: End of second run light map data ....................................................... 112

Table D4: Initial light map data ........................................................................... 113

Table D5: Summary of initial light map data ...................................................... 114

Table D6: GE spectroradiometer data .................................................................. 115

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LIST OF FIGURES

Figure 1: PAR curves based on absorbed light and direct photosynthesis

measurement .......................................................................................................... 18

Figure 2: Cross sectional view of an LED lamp as well as a conventional LED .. 20

Figure 3: Spectral distribution of LEDs versus a dysprosium lamp ...................... 21

Figure 4: High pressure sodium spectrum ............................................................. 22

Figure 5: Diagram of the greenhouse used in the experiment ............................... 32

Figure 6: Light and plant setup for each section .................................................... 36

Figure 7: Total number of marketable fruit per light treatment ............................. 42

Figure 8: Total marketable fruit mass per light treatment ..................................... 43

Figure 9: Total number of fruit per light treatment ................................................ 44

Figure 10: Total fruit mass per light treatment ...................................................... 46

Figure 11: Fruit mass average per light treatment ................................................. 47

Figure 12: Fruit mass average for fruit over 90 g per light treatment .................... 48

Figure 13: Wet mass of plant biomass (excluding fruit) per light treatment ......... 50

Figure 14: Dry mass of plant biomass (excluding fruit) per light treatment ......... 51

Figure 15: Dry to fresh biomass ratio .................................................................... 52

Figure 16: Total fruit mass to plant biomass ratio ................................................. 55

Figure 17: Marketable fruit to plant biomass ratio ................................................ 56

Figure 18: Light direction from the LED arrays .................................................... 68

Figure 19: Typical angular response of the LI-193 ............................................... 70

Figure 20: Two examples of viable clusters turning into secondary stems ........... 77

Figure 21: Leaf burn .............................................................................................. 78

Figure 22: Powdery mildew ................................................................................... 79

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Figure A1: Total marketable fruit for the second run for the tomato plants .......... 89

Figure A2: Total marketable fruit for the first run for the tomato plants.............. .89

Figure A3: Total mass of marketable fruit for the second run ............................... 90

Figure A4: Total mass of marketable fruit for the first run ................................... 90

Figure A5: Total fruit count during the second run ............................................... 91

Figure A6: Total fruit count during the first run .................................................... 91

Figure A7: Total fruit mass during the second run ................................................ 92

Figure A8: Total fruit mass during the first run ..................................................... 92

Figure A9: Total plant fresh biomass (excluding fruit) for the second run ........... 93

Figure A10: Total plant fresh biomass (excluding fruit) for the first run .............. 93

Figure A11: Total plant dried biomass (excluding fruit) for the second run ......... 94

Figure A12: Total plant dried biomass (excluding fruit) for the first run .............. 94

Figure B1: Total number of marketable fruit per plant for the 120 day harvest .... 95

Figure B2: Total marketable fruit mass per plant, for the 120 day harvests .......... 96

Figure B3: Total number of fruit per plant, for the 120 day harvests .................... 96

Figure B4: Total mass of fruit per plant, for the 120 day harvests ........................ 97

Figure B5: Total fresh biomass per plant for the 120 day harvests ....................... 97

Figure B6: Total dry biomass per plant for the 120 day harvests .......................... 98

Figure B7: Ratio of dry to fresh biomass production ............................................ 98

Figure B8: Ratio of fruit mass to fresh biomass for the 120 day data ................... 99

Figure B9: Ratio of marketable fruit mass to fresh biomass for the 120 day ........ 99

Figure B10: Total number of red fruit per plant, for the 120 day experiment ..... 100

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LIST OF EQUATIONS

Equation 1: Number of fruit over 90 grams ........................................................... 40

Equation 2: Mass of fruit over 90 grams................................................................ 41

Equation 3: Number of fruit ................................................................................... 42

Equation 4: Fruit mass ........................................................................................... 43

Equation 5: Vegetative fresh biomass .................................................................... 44

Equation 6: Vegetative dry biomass ...................................................................... 45

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ABBREVIATIONS

ANOVA: Analysis of variance

HPS: High pressure sodium

LED: Light-emitting diode

PAR: Photosynthetically active radiation

R2: Coefficient of determination for statistical model fit

s.e.: Standard error

St. dev.: Standard deviation

5:1: ratio of five times more red light than blue light intensity (μmol m-2 sec-1)

10:1: ratio of ten times more red light than blue light intensity (μmol m-2 sec-1)

19:1: ratio of 19 times more red light than blue light intensity (μmol m-2 sec-1)

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1. INTRODUCTION

Tomato is one of the most consumed crops in the world. In northern climates,

tomatoes have a short time period for growing in the field, thus to maintain supply

and reduce shipping costs, tomatoes are grown locally in greenhouses. Northern

climate greenhouses permit year-round tomato production and provide consumers

with local, competitively priced production. Supplemental lighting is a major

expense but permits increased production, especially during the winter months,

when the daylight interval is much shorter. Greenhouse growers’ benefit from

natural sunlight, and with the addition of supplemental lighting, provide the plants

with a longer irradiance period during the day (usually a 16 hour photoperiod as

opposed to the 10 hours of natural sunlight).

Conventional lighting systems for greenhouses utilize broad-spectrum light

sources, such as high pressure sodium (HPS) or fluorescent lamps. These lamps

are excellent luminous sources for the human eye, but are not the most efficient

light sources for plant production, due to their low levels of blue light and other

photosynthesis sensitive wavelengths. Light-emitting diodes (LEDs) are a fifty

year old technology which is showing potential in the greenhouse industry. With

LEDs, specific wavelengths can be produced, creating a custom light spectrum

targeted for maximum plant production. Studies have shown that the most

important wavelengths for photosynthesis are in the blue and red wavelengths;

peaks in photosynthetic efficiency are found at 440 (blue), 620 (red) and 670 (red)

nm (+/- 10 nm) (McCree 1972a) (Figure 1 in section 3.2).

This research consisted of a full factorial design of three intensities (High: 135

µmol m-2 s-1, Medium: 115 µmol m-2 s-1 and Low: 100 µmol m-2 s-1) and three

ratios of red (661 nm) light to blue (449 nm) light (5:1, 10:1 and 19:1), to

determine which combination was best suited for tomato plant growth and

production. A series of secondary light treatments were added to compare the

factorial results to the current industry standard HPS lighting, a control with no

supplemental lighting, a 50%:50% LED:HPS (10:1 and HPS combined), a 100%

red, and a 100% red where the LEDs were placed at the base of the plants, facing

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upwards. The production increase due to the light treatments was statistically

significant; the top producing treatments (5:1 High and 5:1 Med) were

significantly different from the control (no supplemental light).

Following this introduction is a description of the objectives and basic hypothesis

of the project (Chapter 2). Chapter 3 is a review of pertinent literature which is

used as a basis for the project set-up, as well as a guide of the experimental

design, performance and analysis of the experiment. The methodologies followed

during the experimental phase of this research are outlined in Chapter 4. The

results are presented in Chapter 5, with a discussion of the pertinent quality

assessments of the results as well as an overview of what is needed for future

research presented in Chapter 6. Chapter 7 completes this thesis with an

assessment of the outcome of the research as well as a final conclusion of the

results obtained. Given the industrial nature of the project, specific

recommendations are made for new and existing greenhouses interested in

increasing their production capacity by switching to a partial or full LED

supplemental lighting system.

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2. HYPOTHESIS AND OBJECTIVE

This experiment was conducted within the targeted objectives of General Electric

Lighting Solutions Canada, where the overall goal of the project was to use the

company’s prototype LED arrays to research and determine which ratio and

intensity combinations best suit tomato plants (Solanum lycopersicum), in the

fruiting stage for greenhouse applications. Previous studies performed at McGill

University (unpublished data, Biomass Production Laboratory) have shown that

the ratios of red to blue light used for this experimentation increased plant

photosynthetic rate at the seedling stage, and this research was to confirm if these

same ratios would result in improved biomass production (fruit and vegetative)

for mature plants.

The specific aims were to:

Determine which combination of ratio and intensity of LEDs was

optimum for plant production.

Compare these results to plants grown under HPS lighting, 50%:50%

HPS:LED and a control with no supplemental lighting.

Compare 100% red LEDs in a normal configuration to 100% red LEDs

placed at the bottom of the plants shining upwards.

It was theorized that lighting from below could increase plant uptake of the

energy, thus two treatments were put in place as a side experiment to test this

theory.

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3. LITERATURE REVIEW

3.1. Tomato and Greenhouses

Tomatoes are one of the most important crops in the world. According to

Statistics Canada, tomato sales accounted for close to 50% of the total fruit and

vegetable sales in the country in 2011 (Statistics Canada, 2011). Tomato sales in

Canada reached $496 million, an increase of 5.5% since 2010. The next highest

selling crops in Canada are peppers ($300 million), cucumber ($279 million) and

lettuce ($27.6 million) (Statistics Canada, 2011). Tomatoes can be grown in the

field, but increasingly, tomato production occurs in greenhouses. Greenhouse

tomato production started in the late 1980’s for Canada and the late 1990’s for the

USA, and grew more than tenfold from 1992 to 2003 (Cook et al. 2005).

Greenhouse production exceeded field grown tomatoes in Canada in 1997, and in

2003, the greenhouse production was more than nine times that of field grown

tomatoes (Cook et al. 2005). Today there are 23 million square meters allocated

to greenhouse fruit and vegetable production in Canada, with sales exceeding $1.1

billion annually. According to Brazaitytė et al. (2009), in countries of high

latitude (e.g. Canada), tomatoes are almost exclusively grown in greenhouses.

Light irradiance is the limiting factor for increasing production in greenhouses,

when all other factors (temperature, nutrient levels and water availability) are

adequately maintained (Lefsrud et al. 2008). Ohashi-Kanto (2007) states that

artificial lighting permits stable vegetable crop production no matter the

environmental conditions (with favorable temperature in the greenhouse).

Greenhouses in northern latitudes must compensate for the attenuation in total

light availability (from prolonged winter with short daylight hours), and

supplemental artificial lighting is required in order to maintain a consistent crop

yield throughout the Canadian winters. Conventional greenhouse lighting

systems utilize broad-spectrum light sources, such as high pressure sodium (HPS)

or fluorescent lamps. These lamps were tailored for human vision and therefore

are not ideally suited for plant growth (Bula et al. 1991). For example, light

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irradiance from HPS lights is neither considered optimal nor efficient for plant

growth (Brazaitytė et al. 2009), because it lacks irradiance in the 400-500 nm

range, and this low level of blue light is detrimental to plant growth. Some HPS

lamps have been modified to provide irradiance in the 400-500 nm range (SON-

T-Agro HPS lamp, Philips, Amsterdam), which significantly increased the crop

production when compared to normal HPS lamps (Brazaitytė et al. 2009). These

modified HPS bulbs did not perform as well as most other light treatments tested:

380 nm, 447 nm, 520 nm, 595 nm, 622 nm, 638 nm, 660 nm, 669 nm, 721 nm

LEDs and combinations (Brazaitytė et al. 2009). Modified HPS bulbs can result

in morphological disorders (lower chlorophyll concentrations, smaller stomata

and lower above-ground biomass) in many species, such as tomato plants

(Menard et al. 2006).

3.2. Photosynthesis

Plants require light for photosynthesis, proper growth (seed germination, organ

elongation and phototropism) and physiological development (Goto 2003). Light

is also needed by plants to mediate many hormonal and morphological changes,

and specific wavelengths have been shown to benefit plants in different ways

(Okamoto 1996). For example, both blue light (~420-450 nm) and red light (660

nm) are known to increase chlorophyll production and play an active role in

photosynthesis (Okamoto 1996). Both wavelengths have also been shown to

improve morphological and physiological aspects of plants (Lu et al. 2012;

Poudel et al. 2008). Light catalyzes chemical reactions within the chloroplast

through photosynthesis. Light absorbance occurs in chlorophyll a and b (green

pigments which allow plants to absorb energy from light), as well as in the

carotenoids, lutein and β-carotene (other organic pigments synthesised by plants),

with each pigment having its own absorbance characteristics. Light excites these

pigments which results in improved growth; however, some light wavelengths

which are not absorbed by these pigments can have beneficial effects. For

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example, green light is not absorbed by these pigments, but exposure to only

green light at high photosynthetic photon flux will produce a healthy plant (Lu et

al. 2012).

Figure 1: PAR curves based on absorbed light and direct photosynthesis

measurement (Taiz and Zeiger 1998).

The photosynthetically active radiation (PAR; Figure 1) curve represents the

percentage of light absorbed and utilized by the different pigments in the plant as

a function of wavelength between 400 and 700 nm. The curve has been

developed based on the photosynthetic efficiency (called the action spectrum: the

rate of physiological activity plotted versus light wavelength) and plant pigment

light absorbance curves. Maximum growth occurs in the red and blue spectrums

of light while reduced growth occurs in the green region.

Numerous studies have reported that differences in wavelength produce

significant changes in plant production, photosynthetic efficiency and pigment

accumulation; changes in the irradiance wavelength spectrum have resulted in

changes in biomass production and plant morphology, due to the effect of blue

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light (Hoenecke et al. 1992; Gauthier et al. 1997; Taiz and Zeiger 1998), such as

shorter hypocotyl length (2 mm as opposed to 30 mm), stronger concentration of

chlorophyll (up to 20% higher), while with shifts in the ratio of the red/far red

spectrum (Brown et al. 1995; Heraut-Bron et al. 2001; Hoenecke et al. 1992; Taiz

and Zeiger 1998), there were plant elongation, lower overall dry mass (10% when

compared to white light) and smaller leaf area (15% lower when compared to

white light). Traditional methods of wavelength control (such as filtering white

light, blue enriched white light, or far red enriched white light) can cause changes

in the chlorophyll a to b ratio (Walters and Horton 1995), chlorophyll biosynthesis

and action spectrum (Anstis and Northcot 1974; Björn 1967; French 1991; Koski

et al. 1951; Ogawa et al. 1973; Virgin 1993), and the β-carotene biosynthesis

(Ogawa et al. 1973). Under UV-B light exposure, decreases in chlorophyll b have

occurred (Taiz and Zeiger 1998). Virgin (1993) showed that chlorophyll’s

relative accumulation is wavelength dependent for bean (Pinguicula vulgaris L.),

pea (Pisum sativum L.), potato (Solanum tuberosum L.) and wheat (Triticum

spp.), with both species and individual plant tissues responding differently.

Measurement of the absorption spectra has been determined for most plant

pigments (Köst 1988); limited information exists on the effect of wavelength on

the production of vegetable crops. Existing studies have emphasized the action

spectra of chlorophyll in plants (Anstis and Northcot 1974; Björn 1967; French

1991; Koski et al. 1951; Ogawa et al. 1973; Virgin 1993), but very little work has

concentrated on the action spectra or the effect of wavelength on PAR efficiency,

at intensities above the compensation point. The compensation point is the light

intensity at which the rate of photosynthesis is exactly equal to the rate of

respiration for a plant.

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3.3. Lighting Systems:

Light-emitting diodes (LEDs) were first invented in 1962, by Nick Holonyak

under the name Ga(As1-xPx) p-n Junctions (Holonyak et al. 1962). LEDs are

semi-conductors, which emit light through electroluminescence (when an electric

current or electric field passing through a material triggers a light response).

LEDs are created using a p-n junction, where a p-type semiconductor (positive)

and an n-type semiconductor (negative) are set side by side (Figure 2).

Figure 2: Cross sectional view of an LED lamp as well as a conventional LED

chip (typically 250 x 250 x 250 μm) (Craford 1992)

Different materials give different colours; red light can be a combination of

aluminium gallium arsenide (AlGaAs), gallium arsenide phosphide (GaAsP),

aluminium gallium indium phosphide (AlGaInP), or gallium (III) phosphide

(GaP) (Craford 1992; Mukai 1999). For blue light, Zinc selenide (ZnSe), or

Indium gallium nitride (InGaN) (Craford 1992; Mukai 1999; Xie 1992) can be

used. LEDs have been shown to last up to 100 000 hours, but early degradation in

output or life expectancy will occur in extreme temperatures and high current

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settings (Fu et al. 2011). LEDs life expectancy declines exponentially with

increasing p-n junction temperature (Fu et al. 2011). Conditions of high moisture

and temperature, such as those found in a greenhouse setting may deteriorate the

LEDs faster, but their longevity will still remain higher than that of current

greenhouse lighting technologies (Fu et al. 2011). LEDs emit a short span,

monochromatic light, which permits the creation of a custom light spectrum (by

adding different LEDs), which can more closely resemble the light spectrum

needed by plants for photosynthesis. Figure 3 (below) shows the spectral

distribution of four types of LED lamps used by Xiaoying et al. (2012) with a

dysprosium lamp as a control. The LEDs used were 449 nm (blue), 512 nm

(green), 590 nm (orange) and 632 nm (red). As seen in Figure 3, LEDs promote a

much higher intensity for the wavelengths they emit than a dysprosium lamp

(white light, main difference with HPS is a higher percentage of blue light and

lower red light). Each LED creates a very strong intensity of a targeted

wavelength for photosynthesis, increasing the efficiency. For this experiment, the

LED peaks were centered at 449 nm (blue) and 661 nm (red).

Figure 3: Spectral distribution of different LEDs versus a dysprosium lamp

(Xiaoying et al. 2012)

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High Pressure Sodium (HPS) lights are the major source for greenhouse lighting

(Argus 2010) with over 56 million lamps sold worldwide (Heliospectra 2011).

The use of HPS in greenhouses was shown to significantly increase production in

plants in the early 1970’s, and was shown to have beneficial effects in tomato

production (McAvoy et al. 1984). The increase in plant growth was due to

increased lighting duration in greenhouses and was extended to 18 hours daily (as

opposed to the sole use of sunlight, ~10hr) (McAvoy et al. 1984). Adverse

effects, such as arrested flower development, foliar discolouration or slow growth

of the plant’s apex occurred from the use of HPS lighting on tomato plants

(McAvoy et al. 1984). These adverse effects were explained by the lack of blue

light in the HPS spectrum, which is essential for proper plant growth (Brazaitytė

et al. 2009). The spectral range of HPS lights focuses heavily in the yellow

wavelengths (Figure 4), while lacking red and blue wavelengths, which are

essential for photosynthesis (Figure 1).

Figure 4: High Pressure Sodium Spectrum (Agriculture Solutions 2012)

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3.4. LEDs in Plant Research:

The action of light spectra has been a focus of study since the late 19th century,

but it wasn’t until 1919, with research from Garner and Allard on the

photoperiodic response of flowering in plants that the field of photobiology

developed (Garner et al. 1920). Research has focused on the effect of different

lighting apparatus on plants (Cosgrove et al. 1981; Barro et al. 1989; Brown et al.

1985; Lefsrud et al. 2008; Maruo et al. 2012), and now, LEDs are a major focus

for research in this field. The response of plant parts to LEDs has been researched

since the early 1990’s (Bula et al. 1991; Hoenecke et al. 1992). The rapid

increase in LED technology has been driving this research (Brazaityė et al. 2009).

LED performance has become more than 20 times more efficient from the mid-

1970s to the mid-1990s (Crawford 1992), creating more and more lumens out of

less watts (0.2 lumens per watt in 1970, 10 lumens per watt in 1990). New

colours have been added, such as orange and green lights, but also non-visible

colours such as far-red and ultra-violet (XiaoYing et al. 2011, 2012). Hoenecke et

al. (1992) described the high-output LEDs used for their research as having a

bandwidth of ± 30 nm with a peak at 660 nm, and Xiaoying et al. (2012) reported

new LEDs with a bandwidth of ±15 nm, permitting for much better focus on the

most efficient wavelengths for photosynthesis. This advancement in wavelength

control is not the only benefit LEDs have over other commercially used lighting

sources for plant development: LEDs are characterized by a relative small mass (<

1 g each), small volume, relatively cool emitting temperature, longevity of over

100 000 hours (Folta 2005), linear photon output, wavelength specificity and the

range of possible wavelengths which can be created (Goins et al. 1997; Brazaityė

et al. 2009; Massa et al. 2008) are all characteristics which make them better

suited to crop production than earlier greenhouse lighting systems. Bula et al.

(1991) proposed that the advancement of LEDs would be the solution for growing

plants in space, and that they could potentially be used as a sole light source for

crop growth, as opposed to supplemental lighting. NASA has focused on the

necessary ratio of red light to blue light in order to grow plants in outer space with

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LEDs as their sole source of light, such as for a life-support system on Mars

(Massa et al. 2008). Due to the low availability of blue LEDs, red LEDs were

supplemented with blue fluorescent lights, and the research focused on the

possibility of sole-source or supplemental lighting for outer-space missions, and

controlled environment research (Massa et al. 2008). With the increased

availability of the full range of colors of LEDs and the decreasing costs of

fabrication, LEDs are becoming viable for extensive commercial applications,

such as tomato greenhouses in northern latitude countries; and are being

researched extensively (Brazaitytė et al. 2009). Tennessen et al. (1994) explained

that research on the effect of different wavelengths on plants will greatly benefit

from LEDs since they work as well (and sometimes better, with a quantum yield

of photosynthesis of 0.0027 with red LEDs versus 0.0022 with white light from a

metal halide lamp) as other lighting systems, are more reliable, easily repeatable,

and much more portable.

3.5. Effects of Different Wavelengths

Different wavelengths of the light spectrum have been found to have specific

effects on plant morphology, physiology, photosynthesis efficacy and flowering

capabilities (Menard et al. 2006). Most wavelengths have been shown to have

positive and negative aspects, thus research is focusing more on proper

combinations of light (Massa et al. 2008). Deregibus et al. (1983) showed that red

light (600-700 nm) is beneficial to grasses from the Lolium spp., by increasing the

tiller rate by 20% and increasing the leaf area by 15% without showing any

morphological drawbacks. Red light (650 nm) is shown to have beneficial effects

on tomato and cucumber (Menard et al. 2006). Menard (2006) showed that red

light (650 nm) slightly reduces internode length in cucumber, benefited fruit color

and post-harvest conservation, but can also increase the total amount of starch

molecules inside the chloroplasts in tomatoes by up to 10%.

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Okamoto et al. (1996) reported that both red and blue light can be used by

chlorophyll during photosynthesis, and explained that blue light is beneficial to

plant morphology and overall health: blue light (450 nm) has been shown to

heavily suppress stem elongation in multiple plant species. The effect has been

shown to have effects which can last for many hours into the night (after

termination of the supplemental lighting) in cucumber (Cucumis sativus), pea

(Pisum sativum) and mung bean (Vigna radiata), while plants such as sunflower

(Helianthus annuus), azuki bean (Vigna angularis), and zucchini (Cucurbita

pepo.) undergo dark recovery within the first half hour of night (Cosgrove 1981).

Blue light has shown to decrease dry mass and increase stem length in marigold

seedlings (three times longer stems than under the control or the red supplement),

while it has been shown that the same light would increase dry mass for Salvia

(Salvia spp.) seedlings (Heo et al. 2002). Poudel et al. (2008) showed that blue

light promoted stem growth suppression in grape transplants, but that it also

increased the concentration of chlorophyll in the shoots by 40 to 67% dependant

on the cultivar when compared to red light, making the smaller plants (40%

smaller than when compared to red light) and more effective at photosynthesis.

Overall, blue light is not as effective for photosynthesis as red light, since it

inhibits leaf growth by reducing cell expansion and reduces the total amount of

chlorophyll in the leaves (Goto 2003). Due to this lower efficiency, researchers

tend to undervalue the use of blue light and not consider it in high proportions for

plant growth (Goto 2003). A lack of blue light has been shown to have very

adverse effects on plant morphology: low number of chloroplasts, lower thickness

of cell walls, and low spongy mesophyll tissues (Goto 2003). Blue light was

shown to stimulate stomata opening, and increasing the rate of photosynthesis by

up to 30% in some species (Menard et al. 2006). Growth responses (enhanced

total dry matter, more flowering…) to blue light have been shown to be more

significant and rapid than similar growth responses to red lights (Goto 2003).

Goto (2003) explains that some of the blue light responses are not dependent on

the ratio of blue to red, but just on the total irradiance of blue light. The ratio of

blue to red light is shown to be the most important factor when using LEDs, since

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having both blue light and red light increases plant biomass growth and fruit

production by over 20% when compared to plants grown under only one of the

wavelengths (Goto 2003; Lefsrud et al. 2008; Brazaitytė et al. 2009; XiaoYing et

al. 2011, 2012).

Far red light has been shown to have a strong biological significance, but is not as

readily available on the market as blue and red LEDs. The low availability of far

red LEDs is due to the limited value for human applications or the industrial

sectors (Kubota 2012). Far red has been shown to induce and increase flowering

in many plants such as Gypsophila paniculata and Arabidopsis (Hori et al. 2012),

potato (Miyashita et al. 1995), increase leaf area in lettuce (Kubota et al. 2012),

and stem growth in pepper (Brown et al. 1995). Far red light does not impact

photosynthesis directly, and doesn’t promote dry matter production (Goto 2003).

Supplementing far-red light to red light increases plant growth and health

significantly, by increasing internode length, increasing plant biomass and

increased photosynthesis (Miyashita et al. 1995; Goto 2003). Brown et al. (1995)

suggested that the addition of blue light to red light is much more crucial than the

addition of far-red light to red light.

Plant response to light from the red and the blue spectra has been documented

extensively (Menard et al. 2006). The results show that plant response to LEDs

(and different wavelengths) depends on the plant species and in some cases even

the cultivar. Cosgrove (1981) showed that three different cultivars of cucumber

were not impacted in the same way by the blue light stem development

suppression; Burpee’s Pickler type cucumber suffered from rapid inhibition but

underwent dark recovery (return to normal growth overnight), Levo type pickle

suffered from rapid inhibition and did not undergo dark recovery and finally

Lemon type pickle suffered from the strongest rapid inhibition and remained

stable during the night (Cosgrove 1981).

Other wavelengths have been studied, such as orange and green, showing that

they significantly reduced photosynthesis when compared to other wavelengths in

tomato plants (XiaoYing et al. 2011, 2012). A 200% reduction in photosynthetic

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response was observed for both orange and green wavelengths when compared to

a blue and red light composite, a 120% reduction was observed when compared to

blue light and a 100% reduction was observed when compared to red light

(XiaoYing et al. 2011, 2012). When used in conjunction with blue and red

wavelengths however, green light may have a beneficial effect on overall plant

growth. No statistical differences for production or photosynthetic response were

found between the treatments with only red and blue wavelengths and the

treatment with green light added to the red and blue, but the one with added green

had slightly higher stomatal growth and plant size (XiaoYing et al. 2011). Yellow

and green lights were shown to greatly increase stem length (by over 40% when

compared to white light) and decrease leaf area (by over 50% when compared to

white light) and resulted in the leaves being more brittle and lighter in colour (less

pigments) and lower overall net mass of the plants (XiaoYing et al. 2011).

3.6. Effect of Intensity

Lefsrud et al. (2006) reported that changes in irradiance levels result in significant

changes in the measured accumulation of carotenoids and chlorophylls in kale.

Dorais et al. (1990) showed that an increase in photosynthetic photon flux density

from 100 μmol m-2 s-1 to 150 μmol m-2 s-1 (supplied from 400-W HPS bulbs) will

increase production by 10%, 16% and 14% for tomato plants (Lycopersicon

esculentum Mill. cv. Caruso) grown in low density (2.3 plants m-2), variable

density (between 2.3 plants m-2 and 3.5 plants m-2) and high plant density (3.5

plants m-2) respectively (Dorais et al. 1990).

McAvoy et al. (1984) showed that increasing light irradiance has beneficial

effects on fruit production. Five experiments were run on tomato plants during

the first six months of 1982, using 400-W HPS bulbs to produce 100 μmol m-2 s-1,

125 μmol m-2 s-1 and 150 μmol m-2 s-1. Fruit yield, mass, cluster size and percent

fruit set all increased significantly: average number of fruit increased from 4.4 per

plant (for 100 μmol m-2 s-1) to 5 (for 150 μmol m-2 s-1), mass increased from 2 kg

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per m2 (for 100 μmol m-2 s-1) to 2.9 kg per m2 (for 150 μmol m-2 s-1) (McAvoy et

al. 1984).

A study was done by Tennessen et al. (1994), submitting tomato plants to a range

of intensities from 0 μmol m-2 s-1 to 1400 μmol m-2 s-1, using red LEDs (660 nm)

or water filtered light from a xenon arc. It was show that the photosynthetic

response was similar for the xenon white light and for the red LEDs;

photosynthesis increased until close to 800 μmol m-2 s-1 (with each increase in

intensity causing diminishing increases in photosynthetic rates), and then

remained constant up to 1400 μmol m-2 s-1 (a slight loss occurred with the red

LEDs after 1000 μmol m-2 s-1). Stomatal conductance behaved similarly to the

rate of photosynthesis up to 800 μmol m-2 s-1, but higher intensities decreased the

conductance. It was also shown that the red LEDs promoted higher

photosynthetic rates than the white light until 250 μmol m-2 s-1, where the white

light becomes more efficiently used (Tennessen et al. 1994).

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4. MATERIALS AND METHODS

4.1. Plant Care

All the culturing methods and the tomato care were based on the methods used by

Savoura (Portneuf, QC), a large tomato production greenhouse. The greenhouse

set points were 21oC during the day and 16oC during the night. The watering

system was on for 3 minutes every 20 minutes, starting at 06:00 and ending at

22:00. The supplemental lighting was left on for 16 hours a day, from 06:00 to

22:00. The nutrient solution supplied to the plants via the drip irrigation system

was based on a full strength Hoagland’s solution (Hoagland et al. 1950) modified

by Savoura (proprietary information), of which 40L (nutrient solution) were used

per week (20L of the nitrogen solution and 20L of the micronutrient solution).

The water flow was 40 ml per minute, with total irrigation of approximately 6L

per plant per day (based on 1 ml per 1 J of irradiance, instructed by Savoura). The

nutrient solution was monitored three times a week to maintain proper electro-

conductivity (EC) and pH level. The nutrient solution was mixed automatically,

as needed using one hundred times concentrated stock solutions. Plants were

pollinated by hand, shaking the flowers with a Q-tip.

A central computer in the greenhouse was used as the control system. It

controlled the lighting (on at 06:00 and off at 22:00), the irrigation (on for 3

minutes every 20 minutes during the same hours as the lights), and the ventilation,

coupled with a mister, in order to keep the temperature close to the set points.

The internal greenhouse temperature was monitored and controlled; relative

humidity was monitored, but not controlled.

The LED arrays were positioned at an inter-canopy height, no more than 10 cm

below the top of the plants. The lights height was increased weekly at the start of

the experiment, and every two weeks towards the end, in order to keep the LED

arrays at a level of 75% the plant growth. Once the plants reached the maximum

height of 2.1 m (7 feet) for the greenhouse bays, the array height could not be

increased any more.

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At harvest, fresh mass was determined by separating and weighing both aerial

plant biomass (the rooting system was discarded) and fruit biomass. All fruit

greater than 2 g were counted and weighed (fruit under 2 g were included as plant

biomass). Fruit and flower numbers were counted at two weeks intervals during

the first run, every month for the second run and at final harvest. Fruit was

harvested throughout the experiment at the first observed red pigmentation

(considered ripe), counted and individually weighed. Plants were pruned

according to Savoura’s methods (approximately every 2 weeks), with fresh and

dry biomass measured for each plant (Savoura unpublished data). The Savoura

pruning method was necessary to reduce the leaf nodes to between 10 and 14, to

remove tertiary branching of the stem (a primary and secondary stem were

maintained), but no fruit clusters were removed from the primary or secondary

stems. All fruit were counted (above 2 g); however, only healthy looking flowers

were included. Weekly pruning removed some suckers with flowers: these

flowers were included in the earlier counts, but were not considered in later

counts.

The fresh biomass harvested (aerial and fruit) was dried according to the ASABE

standard (2007), with a temperature of 65 ºC for no less than 72 hours and

subsequently weighed. One representative ripe fruit was collected from each

plant during the final harvests, freeze dried, and stored at -80oC for future fruit

quality measurements. The remaining fruit were made available to Macdonald

Campus students for consumption. No fruit were subject to sensory evaluation.

The plant measurements taken during the experiment included fresh and dry mass,

total and marketable fruit yield, fruit and flower counts, and ratio of fruit to

biomass. Fruit were also separated into two categories depending on size: fruit

from 2 g to 90 g and fruit that weighed more than 90 g. Savoura uses 90 grams as

an internal standard for the minimum mass at which fruit are acceptable to be sold

on the market (Savoura unpublished data).

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4.2. Experimental Setup

4.2.1. Greenhouse Setup

A 7.6 m x 12 m (24’ 10” x 39’ 5”) greenhouse room (set in a north-south

orientation, in the southern west corner of the greenhouse) was used for the

experiment. Long wire-mesh tables (1.2 m high, or 4’) were used as a base for the

tomato plants, in order to ensure more even airflow (with unobstructed areas

underneath the plants. The tables were also set in the north south orientation, with

three tables (4.4 m x 1.6 m, or 14’ 5” x 5’ 4”) in the northern end and three tables

(6.1 m x 1.6 m, or 20’1”x 5’4”) in the southern end of the pod. These tables were

separated into 2 m x 1.6 m (6” 7” x 5’ 4”) sections, which were used for testing.

The southern, longer tables could each accommodate 3 sections, and the northern,

shorter tables could accommodate 2 sections, giving a total of 15 sections for this

experiment. Figure 5 (below) is a map of the greenhouse setup, with the specific

location of each light treatment during the two runs (Table 1, below).

Table 1: Greenhouse Section Placement (with regards to the section numbers in Figure 5, below)

Section Run 1 Run 2 1 5:1 Low 50% LED 2 5:1 Med Red Top 3 5:1 High 10:1 High 4 10:1 Low Red Bot 5 10:1 Med 19:1 High 6 10:1 High 5:1 Med 7 19:1 Low HPS 8 19:1 Med 5:1 Low 9 19:1 High 19:1 Low 10 HPS 19:1 Med 11 Control 10:1 Med 12 Red Top 100% LED 13 Red Bot Control 14 100% LED 10:1 Low 15 50% LED 5:1 High

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Figure 5: Diagram of the greenhouse used in the experiment

Greenhouse Bay with 7 benches (top set) 240 by 64 inch and (bottom set) 173 by 64 inches. Each treatment zone is at least 80 by 64 inches each. Plants were grown in two rows of 4 in a north-south orientation, as shown in zone 11 during the first experiment, and in 2 rows of 3 in a north-south orientation, as shown in zone 13 during the second experiment.

This bay is in the south west corner of the greenhouse with the south and west walls open to the outside environment and the other two walls open to another bay (north wall) or walkway (east wall), the entry door is in the east wall.

Table 1 above lists the placement of each treatment for the first and second experimental run (with respect to the section numbering in this figure)

3

2

1

6

5

4

9

8

7

11

10

Space

not

used

13

12

15

14

N

D

o

o

r

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Each section was surrounded by a double layer of 2.44 m (8 feet) 80% shade cloth

(8MK808, Harnois, St-Thomas, QC) which prevented 96% of light from passing

through, which greatly reduced the cross-contamination of light between

treatments, permitting analysis of all sections independently. A layer of 2.44 m (8

feet) 60% shade cloth (8MK608, Harnois, St-Thomas, QC) was also installed on

top of all the sections, under the roof of the greenhouse. This layer was used in

order to reduce the amount of sunlight attained during the summer (July to

September 2011) to under 20 mol m-2 day-1, in order to simulate winter lighting

conditions. During the second experimental run (January through April 2012),

this layer of 60% shade cloth was removed, in order to get the full winter sunlight.

The amount of natural light received by the plants throughout both experimental

runs was comparable.

A pressure driven drip irrigation system was installed in order to ensure even

distribution of water to the plants. A separate tubing system was built for the

north and the south tables. A 1/2” (5/8” Exterior) black flexible hose (14350

Biofloral, Montreal, QC) was run along the center of each table, and 8 pressure

compensating drip emitters were installed in each section (24 emitters on the

hoses on the tables from the south side, and 16 emitters on the hoses on the tables

from the south side). The emitters used were 19 L h--1 (5 GPH) Antelco agri-drip

emitters (14257 Biofloral, Montreal, QC). Each drip emitter was connected to a

30 cm (1’) 1/4” dripping hose (14314 Biofloral, Montreal, QC) with an 8”

Antelco dripper for 1/4” hose (14259 Biofloral, Montreal, QC), which supplied

water to one plant (set into the rockwool near the stem). The three table length

hoses from each side (North and South) were connected to the greenhouse water

supply. A double tank system was connected to the greenhouse water supply, in

which were stored the two nutrient solutions needed to grow the plants (refer to

section 4.2.)

Tomato plants cv. Trust (esculentum) (grafted onto cv. Maxifort (hirsutum)) were

obtained from Ontario Plant Propagation (St-Thomas, ON), 55 days after seeding.

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Average aerial mass at 55 days was 68.1 g (± s.e. 9.1 g) fresh mass and 4.8 g (±

s.e. 0.7 g) dry mass (10 plants). For the first run, the plants were received as two

single stemmed plants per rockwool cube (5.5 ” x 2.5” x 3.5”) and 4 plants were

placed per rockwool slab (Pargro QuickDrain slab 40”x6”x3” Biofloral, Montreal,

QC). Eight plants (2 rockwool slabs) were placed in each zone, fitted between the

light arrays. For the second run, the plants were received as one double stemmed

plant per rockwool cube (5.5 ” x 2.5” x 3.5”) and 3 plants were placed per

rockwool slab (Pargro QuickDrain slab 40”x6”x3” Biofloral, Montreal, QC). Six

plants (2 rockwool slabs) were placed in each zone. Therefore, there were 8

plants in the first experimental run and 6 plants in the second experimental run.

There were two runs of the experiment: once in summer (July-October 2011) and

once in winter (January-April 2012). A two month period was allotted between

the experimental runs in order to minimize the risk of potential pathogens

carrying over from one experimental run to the other. Each experimental run had

two harvest times: 70 days after being placed in the greenhouse and 120 days after

being placed in the greenhouse. The specific location of each light treatment into

the greenhouse sections was randomly allotted at the beginning of each

experimental run. Half the plants in each section (4 during the first experimental

run and 3 for the second) were randomly selected at the 70 day mark and

harvested, while the remaining plants were grown for the full 120 days.

4.2.1. LED Setup

A full factorial design with two treatment levels was implemented in order to

properly test the different LED ratios and intensities. Three ratios of red to blue

light (5:1, 10:1 and 19:1) and three intensities (Low: ~100 µmol m-2 s-1, Med:

~115 µmol m-2 s-1 and High: ~135 µmol m-2 s-1) were tested. Other experimental

treatments were included to compare the LED treatments to current greenhouse

standard lighting procedures: HPS and lighting from below the plant. The final

fifteen treatments were:

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Table 2: Treatment list and description

5:1 Low 100 µmol m-2 s-1

5:1 Med 115 µmol m-2 s-1

5:1 High 135 µmol m-2 s-1

10:1 Low 100 µmol m-2 s-1

10:1 Med 115 µmol m-2 s-1

10:1 High 135 µmol m-2 s-1

19:1 Low 100 µmol m-2 s-1

19:1 Med 115 µmol m-2 s-1

19:1 High 135 µmol m-2 s-1

100% LED Replicate of the 10:1 High (135 µmol m-2 s-1)

50%:50%

LED:HPS

10:1 Ratio LED coupled with HPS (total: 115 µmol m-2 s-1)

Red Top 100% Red light (130 µmol m-2 s-1)

Red Bot 100% Red light (110 µmol m-2 s-1), LEDs placed at the

bottom of the section, shining upwards into the plant canopy

100% HPS Full HPS light, (105 µmol m-2 s-1)

Control No supplemental lighting

The LED arrays were prototypes from General Electric Lighting Solutions

(Lachine, QC). These consisted of 1.78 m x 8 cm x 2 cm (70 in x 3 in x 0.8 in)

linear fixtures, on which were placed an array of 16 LEDs. A reflective coating

around each LED permitted for better dispersion of the light into the plant canopy.

Each section was fitted with either 3 light fixtures for the Low levels (with the

lights set along the length of the section (Figure 6) or 6 lights (same setup as for

the Low, but having 2 lights side by side in order to increase the intensity). The

LED arrays were designed to provide light at a 45 degree angle in both directions.

They consisted of sixteen modules of three LEDs and a refractor (designed to

project the light in the correct direction), alternating left and right. This way, 8

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groups of LED plus a refractor provide the light to each direction.

Photosynthetically active radiation (PAR) and irradiance levels (W m-2) were

measured to determine light maps of each section, at the beginning of the first

experimental run, the end of the second experimental run and once in between

(see instrumentation sub-section 4.3. for more details).

Figure 6: Light and plant setup for each section

4.3. Instrumentation

Each treatment had a temperature sensor (S-TMB-002; Hobo, Bourne, MA),

located 30 cm below the middle lamp array (except for the 100% HPS and the

Control, where the sensor was placed in the plant canopy at the same height as for

the other sections). Two treatments were randomly selected with

temperature/relative humidity sensors (S-THB-008; Hobo, Bourne, MA), one on

the north side and one on the south side. Light sensors were used in eight selected

treatments using pyranometers (S-LIB-M003; Hobo, Bourne, MA) and quantum

sensors (S-LIA-M003; Hobo, Bourne, MA), recording data for every minute

during the entire experiment. For these eight treatments, a light sensor was

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installed underneath the lamps in the middle of the experimental treatment (to

measure the light from the LED’s) and one was installed above the middle LED

array to measure the sunlight irradiance. For the 100% HPS and the Control, the

light sensor was installed in the plant canopy at the same height as the top sensor

for the other treatment sections. During the first experimental run, light sensors

were placed in the following treatments: 5:1 Low, Med and High, 10:1 High, 19:1

High, HPS, Control, and Red Bottom. During the second experimental run, the

chosen treatments for the light sensors were: 10:1 Low, Med and High, 5:1 Low,

19:1 Low, Red Top, 100% HPS and Control. The environmental measurements

were collected with three data loggers (U30; Hobo, Bourne, MA), and

downloaded to a computer.

A light irradiance map was created by taking nine point measurements beneath

each bulb or array, at a distance of 30 cm (12 inches), 1 m (39 inches) and 1.5 m

(59 inches) from each bulb or array in three locations of each LED array (10 cm

from the ends of the array and the middle location). Two measurement types

were taken for each sensor point: one at a vertical orientation directly below the

bulb and one at a 45º angle relative to the bulb always pointed in a western

direction (at 30 cm toward the plants, first reading measuring 3 arrays, then 2

arrays and one array). This difference in angle was required since the

spectroradiometer measures light in a perpendicular orientation relative to the

sensor and does not include light coming from any other direction; while the

LED’s were designed to direct their light at the plant canopy (to either side of the

lamp at a 45 degree angle). The HPS light data was measured at four levels, top

of the rockwool, starting height of the transplants (1 m, 1.5 m and 30 cm below

the bulb). The second and final light maps were performed with no plants present

in the greenhouse, in order to minimize shade. All readings were taken at least

half an hour after sunset with a spectroradiometer (PS-100, Apogee Instruments,

Logan, UT) with integration of the irradiance to obtain the PAR irradiance values.

A Li-COR spherical quantum sensor was used in the second and the final light

measurements (LI-193; Lincoln, Nebraska), for comparison. Light data are

provided in Table 3 (mid experiment run) and Tables D3-D6.

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Table 3: End of first run and start of second run light map data (January 5th 2012). Compiled data is from average of before (after calibration correction) and end (Li-COR and Spectroradiometer) data

Zone Spectroradiometer Li-COR Average of before and

after measurements

Daily Light Integral from artificial light (mol day-1)

% of Artificial

Light Average

Vertical Average

45º Average Average

5:1 LOW 103.4 135.5 119.4 113.9 98.3 8.50 37.3 5:1 MED 110.2 128.7 119.4 145.8 113.1 9.77 40.6 5:1 HIGH 107.0 136.4 121.7 188.9 132.9 11.48 44.5 10:1 LOW 88.8 121.8 105.3 123.4 104.7 9.05 38.8 10:1 MED 100.1 119.9 110.0 145.6 114.5 9.89 40.9

10:1 HIGH 115.6 140.1 127.8 233.3 151.9

13.13 47.9

19:1 LOW 104.6 139.5 122.0 112.1 104.5 9.03 38.7 19:1 MED 96.8 121.4 109.1 170.6 120.7 10.42 42.2

19:1 HIGH 131.4 154.0 142.7 177.6 139.4

12.04 45.7

LED 131.0 147.4 139.2 204.1 142.3 12.29 46.2 50% 95.2 135.0 115.1 131.3 105.1 9.08 38.8

Red TOP 119.2 149.4 134.3 175.2 133.2 11.50 44.6 Red BOT 82.9 105.6 94.3 98.1 91.2 7.88 35.5 Control 0.0 0.0 0.0 0.2 0.1 0.01 0.1

HPS Light Maximum Irradiance

Middle (1 m) 49.1 49.1 64.7 50.4 4.36 23.4 Height (1.5 m) 91.8 91.8 153.9 106.9 9.24 39.2 Top (30 cm from lamp) 158.7 158.7 427.9 253.9 21.93 60.5

Light map was for the LED bulbs in a vertical and 45 degree angle at a distance of 30 cm from the bulb in 9 locations per section (full darkness, with other bulbs on). The HPS was measured at three heights (of the chamber (approximate height of the seedlings), top of the chamber and 30 cm from the bulb (in a vertical orientation). All measurements were taken with a spectroradiometer and Li-COR spherical quantum sensor.

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4.4. Statistical Analysis

Data was analyzed according to the GLM (generalized linear model; ANOVA)

procedure of SAS (Cary, N.C.), as well as the MIXED procedure. The data was

separated for 70 day and 120 day harvests. The models used involved the Run

factor (experimental run 1 or 2), the Treatment factor (1 through 15, signifying

5:1 Low, 5:1 Med, 5:1 High, 10:1 Low, 10:1 Med, 10:1 High, 19:1 Low, 19:1

Med, 19:1 High, 100% LED, 50% LED, 100% HPS, Red top, Red bottom,

Control, respectively), and the Section Placement in the greenhouse factor

(numbered 1 through 15). The Run and Section Placement factors were treated as

random. A second model was run to make observations about the factorial data

only. The same Run and Section Placement factors were used, but the Treatments

factor was separated into two factors:

- Light ratio (1, 2, 3 for 5:1, 10:1 and 19:1 ratios of red light to blue light)

- Light intensity (100 μmol m-2 s-1, 115 μmol m-2 s-1, 140 μmol m-2 s-1, the

first statistical models were run with 1, 2 and 3, but then switched to the actual

μmol m-2 s-1 values in order to run the regression curves)

The different quantities tested in this experiment were:

- Total marketable fruit number (fruit weighing more than 90 grams)

- Total marketable fruit mass (fruit weighing more than 90 grams)

- Total number of fruit (fruit weighing more than 2 grams)

- Total fruit mass (fruit weighing more than 2 grams)

- Total plant biomass, fresh mass (at harvest)

- Total plant biomass, dry mass (after drying for a minimum of 72 hours,

following the standard ASABE protocol)

- Ratio of fruit mass to fresh plant biomass

- Ratio of marketable fruit to fresh plant biomass (fruit weighing more than

90 grams)

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Based on the statistical method used, plant number was not an independent

variable (since there was only one measurement per plant), thus having a

heterogeneous number of plants for each experimental run (8 plants versus 6

plants) does not impact the statistical analysis. The statistical models used can

handle this heterogeneity.

A standard Tukey-Kramer test was applied to determine significant differences

between treatments. Significance was determined with α ≤ 0.05. The relationship

between experimental dependent variables and treatments was determined by

regression analysis. Orthogonal polynomials were used to study changes

associated with treatments by partitioning the sums of squares into components

associated with linear and quadratic terms. Due to the strong constraints of the

experiment (large plant variability, limited space, limited time, large number of

different factors needing testing), the power of the statistical analysis was

calculated post hoc between 10% and 38% depending on the quantity being

tested.

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5. RESULTS

5.1. Fruit

Fruit data is presented for different categories below. The first category is the

number of marketable fruit, followed by the mass for the marketable fruit. Total

number of fruit and total fruit mass are the subsequent categories, and finally,

average fruit mass is presented. For the factorial model equations below,

quadratic relationships were attempted for all equations, but did not improve the

R2, thus only linear regressions are presented.

5.1.1. Marketable Fruit Number

The top 5 producers of total number of marketable fruit were 50%:50% LED:HPS

(with a total of 118 fruit over 90 grams), Red Top (115 fruit), 5:1 Med (113 fruit),

19:1 High (109 fruit) and 5:1 High (104 fruit). This total data is presented in

Figure 7. For more detailed data see Tables A1 (second run data), A2 (first run

data) and B1 (statistical box plot) in the appendix

Please note that only the factorial portion of the experiment could have regression

equations created, and are explained below.

From the factorial data only, a regression curve with R2 = 0.68 and 4.7 fruit

standard error was fitted. The regression equation was (in number of fruit):

# of Fruit over 90 g = 4.5 + Intensity*0.1 + Light Ratio*(-1.5) (Eqn. 1)

Where Intensity is the amount of light in the treatment (100 μmol m-2 s-1 for Low,

115 μmol m-2 s-1 for Med, 140 μmol m-2 s-1 for High) and Light Ratio is the ratio

of red light to blue light (1 for 5:1, 2 for 10:1 and 3 for 19:1). From this

regression curve, an increase in light intensity promotes a small increase in the

average number of marketable fruit: 1.5 extra marketable fruit when going from

Low intensity to Med intensity, and 2.5 extra fruit from Med intensity to High

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5.1.3. Total Fruit Number

For the total number of all fruit, the highest five producing treatments were 5:1

High (385 fruit). 5:1 Med (358 fruit), 5:1 Low (341 fruit), 19:1 High (315 fruit)

and 100% LED (310 fruit) (Figure A3). Some statistically significant differences

were reported, but the Control was statistically different from most other

treatments, specifically: 5:1 Low, 5:1 Med, 5:1 High, 10:1 High, 19:1 Med, 19:1

High, 100% LED, 50%:50% LED:HPS, Red Top and Red Bottom. This data is

presented in Figure 9. For more detailed data see Tables A5 (second run data),

A6 (first run data) and B3 (statistical box plot) in the appendix.

When analysing the factorial data only, a regression curve with R2 = 0.67 and a

standard error of 6.8 fruit was fitted. The regression equation was (in number of

fruit):

# of Fruit = 23.9 + Intensity*0.12 + Light Ratio*(-4.5) (Eqn. 3)

Where Intensity is the amount of light in the treatment (100 μmol m-2 s-1 for Low,

115 μmol m-2 s-1 for Med, 140 μmol m-2 s-1 for High) and Light Ratio is the ratio

of Red Light to Blue light (1 for 5:1, 2 for 10:1 and 3 for 19:1). From this

regression curve, an increase in Light intensity promotes a small increase in the

average number of total fruit: 1.8 extra marketable fruit when going from Low

intensity to Med intensity, and 3 extra fruit from Med intensity to High intensity

(for an average increase of 4.8 fruit when going from Low to High). An increase

in the percentage of red light decreases the amount of total fruit produced in the

treatment: 4.5 less fruit when going from 5:1 to 10:1 and 4.5 less fruit when going

from 10:1 to 19:1 (for an average decrease of 9 fruit when going from 5:1 to

19:1).

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5.2. Vegetative Biomass

5.2.1. Fresh Mass

Total fresh mass includes the biomass weighed during harvest, as well as the

biomass harvested from the weekly pruning of extra suckers and leaves. The

largest fresh plant biomass was attributed to 19:1 High with 37.8 kg, followed by

Red Bottom with 37.4 kg, 100% LED with 35.3 kg, 50%:50% LED:HPS with

34.5 kg, and 10:1 Med with 33.9 kg. Most light treatments gave very similar

fresh biomass to the plants, with only 3 treatments getting less than 30 kg of

production: Control with 14.7 kg, 5:1 Low with 25.1 kg and HPS with 26.9 kg.

This resulted in an average fresh plant mass between 2700 g for the 19:1 High and

1050 g for the Control. This data is presented in Figure 13. For more detailed

data see Tables A9 (second run data), A10 (first run data) and B5 (statistical box

plot) in the appendix.

When considering the factorial data only, a regression curve with R2 = 0.64 and

standard error of 831.5 g was fitted.

The regression equation was (in grams):

Vegetative Fresh Biomass = 1982.2 + Intensity*5.1+ Light Ratio*426.3 (Eqn. 5)

Where Intensity is the amount of light in the treatment, in μmol m-2 s-1 (100 for

Low, 115 for Med, 140 for High) and Light Ratio is the ratio of Red Light to Blue

light (1 for 5:1, 2 for 10:1 and 3 for 19:1). From this regression curve, an increase

in light intensity promotes a small increase in the plant fresh mass: 76.5 g increase

when going from Low to High, 127.5 g increase when going from Med to High

(for an average increase of 204 g from Low to High). An increase in the

percentage of red light significantly increases the amount of plant fresh mass

produced in the treatment: an extra 426.3 g is produced on average when going

from 5:1 to 10:1, and an extra 426.3 g is produced on average when going from

10:1 to 19:1 (average increase of 852.6 g when going from 5:1 to 19:1).

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5.2.3. Dry to Fresh Biomass Ratio

A dry mass to fresh mass ratio was tested, for the full data set. No statistical

differences were measured, with Control being the lowest (6.0%) and 5:1 Low

being the highest (7.8%). This ratio is used to determine if the plants become

more woody or retain more water (succulent) focusing on the generated biomass

(Clifford 1987). According to the literature, the average value for tomato plants is

between 6 to 8% (Heuvelink 2005). The top 5 treatments for dry to fresh biomass

ratio were 5:1 Low (7.8%), 5:1 High (7.6%), 10:1 High (7.6%), 19:1 High (7.4%)

and 19:1 Med (7.4%). Data can be found in Figure 15, with a supplemental

statistical bar graph in Figure B10.

Figure 15: Dry to fresh biomass ratio No statistically significant differences were observed.

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5.2.4. Fruit and Flower Counts

The fruit and flower counts occurred on August 2nd, 9th, 12th, 16th, 25th, Sept

1st and 28th for the first run of the experiment, and on Feb. 8th, 28th, and April

27th for the second run (Table 4 with additional data in Table C5, appendix). The

5:1 ratio treatment had the most flowering and fruiting, followed by the 10:1, then

19:1 treatments, similarly to the first run. The 5:1 and 10:1 ratios, with middle or

high intensity had the best fruiting results. The 5:1 High had the highest fruit

count (191 fruit in the first run and 194 fruit in the second run), followed by 5:1

Med (192 fruit in the first run and 166 fruit in the second run), 5:1 Low (173 fruit

in the first run and 168 in the second run) and 10:1 High (161 fruit for the first run

and 143 for the second run). Control was the lowest with 13 fruit in the first run

and 10 fruit in the second, followed by HPS with 53 fruit in the first run and 96 in

the second.

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Table 4: Summary fruit and flower counts for the tomato plants (8 per zone for the first replication until September 1st and 4 after, 6 per zone for the second replication)

Date 5:1 TOTAL 10:1 TOTAL 19:1 TOTAL Run 1 Flower Fruit Flower Fruit Flower Fruit 2-Aug 78 61 106 43 102 42 9-Aug 161 85 153 61 118 55 12-Aug 163 116 113 106 85 75 16-Aug 114 169 77 104 63 90 25-Aug 114 277 79 214 71 148 1-Sep 70 411 53 286 41 206 28-Sep 54 205 54 137 54 95 Run 2 8-Feb 58 296 56 294 44 221 28-Feb 300 285 227 248 214 229 27-Apr 316 394 664 280 521 318

LOW TOTAL MED TOTAL HIGH TOTAL

Run 1 Flower Fruit Flower Fruit Flower Fruit 2-Aug 106 15 99 88 81 43 9-Aug 107 27 148 103 177 71 12-Aug 119 40 112 145 130 112 16-Aug 73 59 86 146 95 158 25-Aug 96 152 76 218 92 269 1-Sep 71 192 33 323 60 388 28-Sep 42 145 56 126 64 166 Run 2 8-Feb 33 261 37 235 88 315 28-Feb 206 219 211 238 324 305 27-Apr 452 311 617 315 432 366

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5.4. Environmental Data

5.4.1. Temperature

The greenhouse temperature from December 31st to April 27th ranged from 40.2

ºC to 0.9 ºC. These extreme values occurred due to the greenhouse climate

control system shutting down twice during the month of February, and once in the

months of March and April. The extremes only occurred for a few minutes for

the heat, and a couple of hours for the cold. The average temperature was 16.9 ºC

with a standard deviation of 3.4 ºC, and was very consistent throughout the

greenhouse. The values broken down per month and per section can be found in

Table D2 for the first run and Table D1 for the second run. For the first run, the

average temperature was 21.8ºC with a standard deviation of 2.4ºC. These

conditions are similar to the conditions in a commercial tomato production

greenhouse with short extremes (38.7ºC and 9.9ºC for the highs and lows).

5.4.2. Relative Humidity

The relative humidity was measured in three locations with less than 2%

difference between the sensors. The relative humidity profile was very similar

during both runs of the experiment. Relative humidity had an average of 63%,

with a range from 94.5% to 13.5%. Typically, the average daily highest humidity

recorded was 82% +/- 5% with an average low of 30% +/- 10%.

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5.4.3. Light Sensor Map

The light sensor data is presented in summary and as mapped averages (Table 3,

section 4.3 and tables D3-D6, appendix). The spectroradiometer was recalibrated

for the data in Table D3, with the correction factor used in Table D5. An LI-193

Spherical Quantum Sensor collected light data and is presented in Tables D3-D4.

A spectroradiometer was used from GE and this data is presented in Table D6, but

due to the very large values obtained from this device it was not used for any

analysis (peak values of over 1600 μmol m-2 s-1 were obtained, making it close to

10 times higher than the expected values, or the other sensor recordings). The

large fluctuations in intensity observed with this instrument made it unreliable for

the purpose of the experiment. The data used in the calculations is the combined

data from the spectroradiometer and spherical light sensor taken between both

experimental runs, since those two sensors were found to be the most reliable and

stable for LED readings in this experiment. The first light data (before the

experiment) is reported but due to variation between the different sensors was not

accepted or used in the calculations. The remaining irradiance levels (end of first

run and end of second run) were very similar, with Low intensities having ~ 100

μmol m-2 s-1 (average of 102.5 μmol m-2 s-1 and 101 μmol m-2 s-1 for the second

and last light maps, respectively), while the Med had a level of 115 μmol m-2 s-1

(average of 116.1 μmol m-2 s-1 and 115.9 μmol m-2 s-1 for the second and last light

maps, respectively) and the High had a level of 135 μmol m-2 s-1 (average of 141.4

μmol m-2 s-1 and 130 μmol m-2 s-1 for the second and last light maps, respectively).

The HPS irradiance level average in the chamber was 110 μmol m-2 s-1 (average

of 106.9 μmol m-2 s-1 and 113.2 μmol m-2 s-1 for the second and last light maps,

respectively).

The experiment monitored the natural light irradiance and artificial light provided

to the plant through a number of sensors within the plant canopy, above the plant

canopy below the lights, and above the lights below the shade cloth. The

maximum natural irradiance measured in the test area was at 513 μmol m-2 s-1 at

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noon, which is very similar to the maximum of 566 μmol m-2 s-1 measured during

the summer under the shade cloth. During the summer run, all treatments

received almost equal amounts of natural light (small differences in shading using

the different lighting arrays), but during the winter run, it is possible that the angle

of incidence of the sun had an impact on the distribution of the sunlight. The total

light provided by natural light was an average of 14.3 mol day-1 with a standard

deviation of 6.2 mol day-1. Artificial light provided to the treatments varied from

8.5 mol day-1 for 5:1 low to a high of 13.1 mol day-1 for 10:1 high, the HPS

treatment was at 9.2 mol day-1 (Table D3). The inclusion of the natural light

increased the average daily light integral to an average from 26.3 mol day-1 for

19:1 High, a low of 22.8 mol day-1 for 5:1 Low, and 23.5 mol day-1 for HPS.

These results show that the percentage of total irradiance (including sunlight and

LED light) provided to the plant as artificial light sit in between 37% for the 5:1

Low treatment and 47.9% for the 10:1 High (Table 3, section 4.3). The HPS

treatment was at 39.2%, and the Control was around 0.1%. Irradiance levels

measured within the plant canopy at 1.5 m from the rockwool cubes (bottom of

the leaves on the plants), averaged less than 2 mol day-1.

Table 3 (section 4.3) shows the light map taken during the time in between both

experimental runs. Table 3 data was used for all calculations of light levels for

the experiment, such as the regression equations. Data from the final light map is

presented in Table D3. Table D4 shows the first light map, which was done only

with the spectroradiometer. Table D5 takes the data from D4 and applies the

correction factor sent to us by the manufacturer for the calibration before the final

light map. Little differences were found between Table 3 and tables D3 and D4,

making the use of Table 3 data for calculations viable.

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6. DISCUSSION

6.1. Ratios chosen

The primary focus of this experiment was to test the effect of the LED ratio and

light intensity (the 3x3 factorial). The 5:1, 10:1 and 19:1 ratios were chosen from

previous research performed at McGill University (unpublished data, Biomass

Production Laboratory), which showed that these ratios resulted in improved

photosynthetic responses with plants at the seedling (3-5 true leaves) plant age.

The three intensity levels were chosen following the supplementary light levels

used by Savoura in their greenhouses.

A range of factors or comparison tests were included:

HPS (a standard greenhouse light in order to compare the LEDs to

currently used technology)

50%:50% LED:HPS (a mix of HPS and 10:1 LEDs)

Control (no supplemental lighting, only sunlight)

100% Red Top (can be used to compare with the main tests, to illustrate

the need for blue light, but was installed as a side experiment, to compare

to the 100% bottom)

100% Red Bottom (used as a side experiment, in order to see the effects of

supplying the light from below the leaves of the plants, had an effect on

production)

100% LED (10:1 ratio, meant to be at a higher intensity than the High, at

160 µmol m-2 s-1). Due to the high heat emanating from the fixtures at this

intensity, the biomass which came in contact with the leaves would

experience heat blanching and stop growing. Therefore, the intensity was

lowered to 135 µmol m-2 s-1, making it a replicate of the 10:1 High

treatment.

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Most of the earlier work examining the impact of wavelength occurred under

ultra-low irradiance levels (<13 μmol m-2 s-1) (McCree 1971, 1972a; Inada 1976;

Sager et al. 1982; and Ogawa et al. 1973). Providing plants with elevated

intensities of light at narrow wavelengths will provide a better understanding of

how they utilize these wavelengths. Given that pigments absorb light at specific

wavelengths, conditions should exist whereby the wavelength intensity reaches a

level that result in the optimum light absorbance or possibly, the degradation of

plant pigments. Knowledge concerning the potential range of wavelength

intensities will promote more accurate design of plant lighting systems and

procedures, maximizing plant production

6.2. Fruit

6.2.1 Marketable Fruit

It is important to note that the majority of the fruit harvested were not fully

mature, at the 70 day and the 120 day mark. Thus, the large majority of the fruit

considered non-marketable (< 90 g) would continue growing in a normal

industrial setting, and reach the cut-off (since they are not harvested on a specific

date but after they have matured). Note that the ripe fruit were almost exclusively

considered marketable fruit, since they were harvested when mature. Therefore,

in order to get a proper overview of the effectiveness of a treatment, the results

cannot be based solely on the marketable fruit quantities; total fruit number and

mass also play a large role in the effectiveness of a treatment within this study.

Coupled with the shade cloth used to prevent light contamination from one

treatment to another which blocked out a portion of the sunlight, it results in a

lower overall production than what would be expected in a commercial

greenhouse using the LED lighting system.

The highest producing treatments of marketable fruit were 50%:50% LED:HPS,

5:1 High and 19:1 High . The 100% red treatments (both the top and the bottom)

were ranked fourth and fifth. The 19:1 High, and both 100% red treatments were

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much stronger with the production of total marketable fruit during the second run

when compared with the first run. This contradicts what the literature says about

red light, which is that it promotes biomass production and doesn`t promote

fruiting as much as blue light (Brown et al. 1995, Hoenecke et al. 1992). It is

possible that due to the increased biomass production, these treatments grew

faster during the winter (where the sun was weaker and at more of an angle), and

reached the top of the shade cloth (where more sunlight was available) quicker,

resulting in more production than the treatments which grew slightly slower. It

was also theorized that it could be due to how these treatments were placed in the

south-eastern side of the experiment, closest to the windows (Figure 5), which

benefited from more direct sunlight (with the angle of incidence and lower

irradiance due to the winter, could make a difference). 50%:50% LED:HPS had

the highest marketable fruit production during the second run, and was located in

the south-eastern corner, which started the idea that the location might be a factor

of influence in the study; however 50%:50% LED:HPS was rated second best

(behind 5:1 Med) during the first run, where it was located at the exact opposite

corner of the greenhouse. Angle of incidence of the sun was shown to be not

significant in the statistical analysis. Being ranked in the top 2 for total

marketable fruit production in both runs shows that 50%:50% LED:HPS is one of

the top treatments for marketable fruit. This agrees with research from Menard et

al. (2006) which shows that supplementing blue and red light to HPS creates

much higher production than with HPS alone. The 5:1 ratio had the highest

overall ratio, with all three intensities ranking in the top 50% of the experiment

for both experimental runs. This result is supported by the literature, where blue

light promotes fruiting and flowering (Menard et al. 2006, Goto 2003), but

contradicts Miyashita et al. (1995) who reported that red light, and not blue light,

would increase flowering.

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6.2.2. Total Fruit

The lower limit for considering a fruit was 2 g. This was selected because fruit

mass under this level are hard to distinguish from aborted flowers or aborted fruit;

during the first experiment especially, many small fruit and flowers were aborted

due to the extreme heat and a cut off was needed in order to remove them as

viable fruit. Unfortunately, due to the size cut off, a certain number of viable fruit

have not been considered (especially during the second run, where less fruit were

aborted since less high temperature extremes were felt). These small viable fruit

would have grown into full fruit if the experiment had been allowed to run for

longer, or in a production greenhouse.

The highest producing treatments were 5:1 High, 5:1 Med, 100% Red top,

50%:50% LED:HPS and 5:1 Low. As expected from the literature, higher

intensities bring forth more production (McAvoy et al.1984, Tennessen et al.

1994), with all ratios producing more in the High than the Med or the Low

irradiance levels. The 5:1 ratio was the top ratio from this experiment, with all

three intensities ranking in the top 5 treatments. This agrees with research from

Menard et al. (2006) and Goto (2003), who found that blue light promoted

flowering and fruiting.

The 19:1 High treatment resulted in much higher fruit production for the second

run than for the first run. This may be because 19:1 High had an increased

biomass production from the increased red light, as shown by Brown et al. (1995)

and Hoenecke et al. (1992). This higher biomass could have allowed the plants to

reach the natural sunlight faster than the other plants (due to the side wall

screening lowering the total light incoming from the sun). This extra production

was also observed in both the 100% Red treatments (top and bottom), but not to

the same extent. An increase in biomass from red LED light has been reported

(Brown et al. 1995; Hoenecke et al. 1992). This increase in the earlier stage

allowed for more sunlight towards the end of the experiment (since the plants

could grow taller, they received unobstructed sunlight quicker, increasing the

overall fruit production). This would not have occurred in an industrial

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greenhouse setting, since there would be no shade cloth impeding the sunlight on

the plants, and all treatments would have benefited from maximum sunlight from

the start. Second, due to the placement of the 19:1 High during the second run (in

the middle), the plants were less impacted by the cold (Adams et al. 2001), due to

a larger distance from the windows (data shows that the minima in this section is

over one degree higher than in sections closer to the windows). From the

temperature data however, the 100% LED (10:1 ratio) had a very similar

temperature to 19:1 High (and was also in the more sheltered spots), but did not

perform as well. Thus it is assumed that the red light was the driving factor of the

increased production (Barro et al. 1989), not temperature, which caused early

stem elongation leading to taller plants and more overall light (height was not

measured in the plants during this experiment).

Fruit quality, taste and color were not monitored or tested during the experiment.

No observable differences were noticed in these traits between the treatments

during or after the experiment upon consumption of the fruit.

6.3. Vegetative Biomass

The plants require a minimum amount of leaves to maximize fruit production but

extra vegetative production was pruned in order to maximize yield, since extra

suckers and leaves drain the plant of energy which could be focused into the fruit.

Pruning was performed every couple of weeks, and the biomass from the prunings

was weighed fresh and dry, and added into the final biomass data. The fresh and

dry biomass had very similar patterns to each other; the 5:1 ratio did have the

highest dry to fresh ratio, which coincides with data found in the literature,

showing that blue light promotes an increase in dry mass in tomato plants

(Menard et al. 2006).

The top treatments for both fresh biomass and dry biomass were 19:1 High and

Red Bottom, with the other 19:1 treatments and the 100% Red top in the top 50%

of treatments for biomass. This follows the literature which shows that red light

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produces more biomass (Brown et al. 1995; Hoenecke et al. 1992), since all

treatments with high red light were found consistently towards the top.

100% LED, 50%:50% LED:HPS and 10:1 Med were in the top 5 treatments,

which contradicts what was seen by McAvoy et al. (1984) and Dorais et al.

(1990), where an increase in intensity promotes an increase in biomass. All of

those treatments use the 10:1 ratio, yet the 10:1 High was behind both the

50%:50% LED:HPS and the 10:1 Med, which had 115 μmol m-2 s-1 as opposed to

the 140 μmol m-2 s-1 for the high. 100% LED is a replicate of the 10:1 High and

had increased biomass when compared to the two mid-level treatments which

could suggest that the 10:1 High treatment produced lower than expected due to

plant variability, which would then confirm the research which showed that

intensity had a positive effect on plant biomass (Dorias et al. 1990; McAvoy et al.

1984). 100% Red bottom produced more than 100% Red top, goes against the

research (since the red top received 130 μmol m-2 s-1, as opposed to the 110 μmol

m-2 s-1 received by the red bottom), resulting in less intensity creating higher

biomass. This would suggest that illuminating plants from the bottom could

increase overall plant growth. Moss (1964) showed that illumination from below

was at least as efficient as illumination from above; this research suggests that

illumination from below could increase biomass production significantly.

Ideally, all flowers will become fruit, but some flower abortion, as well as fruit

abortion was noted. The elevated temperature at the end of July, along with the

cold temperatures in February both could have limited the fruit set. Fruit and

flower counts are typically not used to make decisions in selecting significant and

non-significant treatment effects but are used to provide support for the decision

from the yield data (marketable and total).

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6.4. HPS to LED comparison

One major difference between the HPS lighting and the LED lighting is that the

HPS were fixed at the top, to mimic conventional greenhouse setup, while the

LEDs were set at an inter-canopy height. Thus, the LED illuminated plants

benefited from a higher light intensity at the start of the run, while the HPS plants

had less light to begin the experiment. Once the plants grew past a height of ~1.5

m, the HPS illuminated plants received more light than the LED treatments. This

increased light could explain that the LED treatments consistently outperformed

the HPS in every category; this is a benefit inherent to the design of the LED

systems, which can be placed closer to the plants with much less risk of damaging

or burning them (Brazaitytė 2009). Inter-canopy lighting is impossible with HPS

lighting due to the heat emitted by the lamps (Hogewoning et al. 2007).

6.5. Top versus Bottom Lighting Systems

The Red Top and the Red Bottom treatments were directly compared to each

other. The major difference between the two treatments was the set-up with the

bottom treatment lights placed at the bottom of the section and shone upwards,

while the red top LEDs were placed at the top, similarly to all other treatments.

The Red Bottom had slightly less light levels than the Red Top. Overall, the Red

Bottom section slightly underperformed when compared to the Red Top in

fruiting categories, but created more biomass. No statistical differences were

measured between these two treatments. The differences between the two

treatments are more pronounced towards the end of the run. This could be

explained by the lower light levels or because the plants in the Red Bottom

section were growing more side stems and suckers very close to the LED arrays,

and did not grow as tall (observed but not measured). This slight difference in

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yield between the two treatments observed towards the end of the run could be

explained by the Red Top plants growing taller more quickly, reaching the sun

(above the shade cloth) and benefiting from the added natural sunlight quicker.

While, the Red Bottom plants were observed as more bushy and shorter. No

conclusions can be drawn from the statistics, but the production results were

similar for both treatments, and the biomass production was higher in the Red

Bottom despite the light intensity difference. This could suggest that the light

coming from below was at least as beneficial as the light coming from above

(Moss 1964), and possibly even more beneficial than traditional lighting from

above.

6.6. Light Measurement Techniques

The point source nature of the LED arrays makes it very difficult to measure the

light intensity with conventional light measuring devices. Typical

spectroradiometers and light measuring devices are built to measure light levels

from the sun or other light sources which disperse light in every direction (such as

HPS lighting), and therefore are not as compatible with the LED arrays. A series

of irradiance tests were performed with a range of different light measuring

devices, in order to find the most appropriate solution for measuring the LED

point source light and arrays. The first test was performed using a pyranometer

(total solar radiation) (MP-100, Apogee Instruments, Logan UT) and a quantum

meter (measures only the PAR) (MQ-100, Apogee Instruments, Logan UT). It

was found that the light measurement from both devices was very variable (30%

differences in readings), under weighing blue light and over weighing red light

(Apogee 2012a, 2012b), and therefore they could not be used for the experiment.

The light fixtures were created to supply most of their light irradiance at a 45o

angle, as shown below.

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The light measurements were taken at horizontal, vertical and 45o, at three

different heights, at 3 points along the length of each light fixture, in order to get a

better overview of the light received by the plants. A base was created to place

the light measuring device at the exact same distance and angle from the lamp in

each position, but it was found that the measurements were still very variable

(close to 25% variability in the readings). The way the light fixtures are created,

LEDs are facing alternatively to the right or to the left (in order to get the 45o

angle of incidence, the LEDs were not designed to face down), with a reflecting

cone around each LED for better dispersion. Therefore, on the other side of the

LED, the light levels decrease dramatically. The same occurs for light

measurements from the bottom. Due to the reflecting cones, the light level

difference from being exactly under one of the LEDs or a couple centimeters off

makes a significant difference (60% of total radiation) in the light reading. Since

the light measuring devices only measure light coming directly from above, it

does not take into account the light originating at an angle from the LEDs

adjacent to the one sitting directly above the light measuring device (Figure 18

depicts the light direction from the LED arrays, showing that most of the light

does not go downwards).

Figure 18: Light direction from the LED arrays. Longer distance shows higher

light levels.

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Due to the relative simplicity of the quantum meter and the pyranometer, a more

complex spectroradiometer (BLACK-Comet Concave Grating Spectrometer,

StellarNet Inc., Tampa, FL) was tested, with the same measurement positions

used. Previously, the Apogee Instruments light sensor provided a value of 30

μmol m-2 s-1 for the 10:1 Low (average), while the spectroradiometer provided a

value of 68.4 μmol m-2 s-1; showing that the hand held sensor recorded half of the

expected value. Since the hand-held sensors (Apogee Instruments, MP-100 and

MQ-100, Logan, UT) were calibrated for solar irradiation (Apogee 2012a, 2012b)

they may not properly report wavelengths outside the 460-660 nm range. The

spectroradiometer was much more reliable than the quantum meter and

pyranometer, but the same problem with variability due to spatial position was

found. The spectroradiometer was designed for conventional overhead light

sources, and only records light incoming from directly above the sensor. The

field of view of a typical spectroradiometer was shown to be 10o (Mac Arthur et

al. 2007). With adjustments to the positioning (directly facing the LED for the

measurements), a light map could be created using the spectroradiometer, but it

was still not adequate for proper measurements of the light received by the plants.

Please note in Table D6 a spectroradiometer provided by General Electric (GE)

was used; the values have been provided but were significantly higher than any of

the other measurements taken with other instruments and were excluded from the

data analysis. The instrument was much more unreliable, with peaks at 1600

1600 μmol m-2 s-1, while some readings were lower than the expected values. Due

to this large variability, this instrument was not used for calculations. Note that

having a light level above 1600 μmol m-2 s-1 is considered brighter than the sun

and over 10 times higher than what was expected from the greenhouse lighting

system..

An underwater spherical quantum sensor (LI-193, Li-COR, Lincoln, NE) was

tested. The spherical quantum sensor was developed to better understand the light

dispersion in underwater biological experiments, by measuring the photon flux

coming from all directions. This sensor was tested due to its ability to record the

photon flux coming from all directions, which was better suited for measuring the

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LED lighting in this experiment (since the LED arrays do not give out uniform

light). The typical angular response to light is shown below (Figure 19). It is

seen that some light loss occurs, mainly due to the sensor at the base of the

sphere, but that the reading of the light coming from the upper 180o comes with

no significant loss. The same sensor positions were used to take the

measurements (3 heights, 3 positions along the length of each lamp, horizontal,

vertical and 45o). This sensor was well adapted to the LED fixtures, and gave us

much more accurate measurements of the lighting. This was the first sensor

which was found to accurately measure the light received by the plants, since it

took into account the light coming from multiple LEDs at each spot. Some losses

were found: when the bulb is placed sideways, the light measurement was 20%

lower than when placed right side up, which can be explained by the loss of

efficiency in the angular response graph. The Li-COR underwater quantum

sensor was chosen as the most reliable sensor for this research. It was also shown

to be the most adapted sensor for measuring light from LED light arrays available

on the market.

Figure 19: Typical angular response of the LI-193 (Li-COR 2012)

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Both sensors used for the light testing have some limitations: the

spectroradiometer’s angle of incidence is extremely narrow, giving unstable

readings from the point source LED’s, and the spherical Li-COR may have

limitations in the spectral response curves. Provided in this report is an average of

all of this data recorded in order to try and reduce the error that might occur from

this variability. The spectroradiometer had been calibrated before taking these

readings.

6.7. Production Issues

A certain number of production issues occurred during both experiments, possibly

having a slight effect on the overall performance of the plants in the experiment,

and have been outlined below. The production issues were separated per

experimental run.

Almost of all these production issues occurred in a fashion where they impacted

all treatments equally, thus they did not impact the statistical comparison between

treatments. However, some impacts may have resulted in a difference in absolute

values between the results from this experiment and what would be expected from

a commercial greenhouse. The exceptions which could have resulted in statistical

differences since they didn’t impact all treatments were: the higher temperature of

the bulbs for the Med ratio test treatment, the lamps failing (Red bottom, 19:1

Low and 10:1 Low), the lower irradiance level for the HPS treatment, the

increased powdery mildew on 5:1 Med (only in the first run of the experiment),

and edge effects from solar loads and the lamp falling and crushing a plant in the

19:1 Med. The lamp failures did not impact the results statistically. It is unsure

how high of an impact the higher bulb temperature (Med level) or the lower

irradiance in the early stages of the HPS has had on the final results but based on

the statistical models, no major statistical impact was noticed. The results of the

plants showed a larger difference than this lower light level would explain.

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6.7.1. First Run:

The plants were received as two plants per rockwool cube in the first run, which

was not as expected. Changes in irrigation and original planned spacing had to be

put in place, which meant that the plants had to be irrigated manually. The

computer control system was not automated at first, resulting in this manual

irrigation taking a week longer than expected. No impact was experienced in the

statistics, since all the plants experienced this similarly. Once the automatic

system was running, it took four days in order to get the correct amount of

irrigation (Savoura internal standard). Overwatering occurred during this period,

but due to the quick draining nature of the rockwool, excess water was flushed.

No impact was experienced, since all plants experienced this in the same fashion.

The nutrient solution supplied to the plants was in the wrong ratio (much higher in

iron and magnesium and lower in nitrates) for the first 4 weeks of the experiment.

No impact was noticed in the statistical analysis, since all plants were impacted in

the same manner. This could however partly explain why the total fruit mass

produced during this run was slightly lower than during the second run (no

statistical differences). A nutrient solution stronger in nitrogen during the early

stages of plant growth could increase plant mass by close to 50% (Ma et al. 2006).

The first pruning was delayed during the first two weeks, allowing for increased

biomass, flower and fruit counts. A secondary stem was kept in order to limit the

loss of fruiting, but the extra energy used for the second stem could lower the total

fruit production. All plants experienced the same, hence no impact was measured

in the statistics.

The red bottom LEDs stopped working twice due to water vapour condensing in

the housing (corrected after a week the first time and five days the next). Since

only one of the three lamps stopped working and the problem was fixed relatively

quickly, no impact was noticed. Some lamps became unplugged (the weight of

the lamps pulled down on the plugs at the beginning of the experiment) for a

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maximum of 2 days, but were fixed within a few hours, with no measurable

differences. 19:1 Low and 10:1 Low bulbs suffered from failures, and one light in

each section was off for 3 days and 7 days, respectively.

The LED systems originally used in the Medium intensity level (115 μmol m-2 s-1)

were shown to heat up to 88oC (due to the heat generated from the electric box

powering the LEDs as well as the heat accumulated from the sunlight on the black

casing). Since the lamps were set in an intercanopy setting, the leaves and stems

closest to the lamps experienced burning and blanching. As described by

Hogewoning et al. (2007), heat over 40oC close to leaves and stems will burn

them, lowering productivity. This would have lowered the total production from

the plants, since the burnt organs (stems or leaves) lost part of their functionality

(resulting in stunted growth or poor photosynthesis). This was corrected after 3

weeks, by using the same type of system as for the High intensity treatments (2

lamps side by side) which does not generate as much heat. No stems were

impacted and no statistical effect was measured. Heat waves during the summer,

where maximum temperatures were over 30ºC (for a period of 2 weeks, max

recorded was 38ºC) could be responsible for the flower abortion observed. The

heat of the summer has been shown to cause abortion of the flowers and fruit in

tomato plants (Sato et al. 2001). This abortion was observed for all plants in the

greenhouse, and could have an (<10%) impact on the total production from the

first run when compared to the second run. During the peak of the summer, the

fog system failed for 2 days, preventing the main cooling mechanism from

working. One of the fans also failed, and had to be replaced.

Powdery mildew was noticed on some plants, mostly in the 5:1 Med intensity

section. The powdery mildew originated in the 5:1 Med treatment, and slowly

contaminated all treatments in the greenhouse. The occurrence of powdery

mildew in a resistant strain of tomato plants such as the one used for this

experiment is rare, but can occur from exposure (Jones et al. 2001). Up to 25%

decrease in yield potential is possible if the plants are not treated, and the fruit

mass and volume can be decreased by up to 30% (Jones et al. 2001). This could

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explain why the 5:1 Med treatment had smaller fruit than the other top treatments

during the first run.

The measure of irradiance from the LEDs changed over time of output.

Corrections were made every two weeks, in order to make sure the voltage stayed

in the correct range. It was observed that one lamp in the 10:1 High lost voltage

(decreased light intensity) much more rapidly than others, and had to be checked

more frequently.

The HPS intensity was lower than expected at first. After 6 weeks this was

corrected, by adding a second lamp. A decrease in yield from the HPS plants is

expected from this lower intensity, but a conjecture was made for correction

(accounting for a 50% loss in production, which is higher than the loss

calculated). The solar light irradiance at different location in the greenhouse was

different. This has a low impact during the summer (when the sunlight was

directly overhead) but may have an impact during the late fall (when the sun was

lower). The southern treatments (5:1 low, 10:1 low and 19:1 low) could have

received more solar irradiance, as well as the eastern treatments (all 5:1

treatments, control and HPS). The difference in production is not expected to be

significant, and since the top treatments continued producing better during the

second run despite less favourable locations, the difference of solar light

irradiance was assumed to be not significant.

6.7.2. Second Run:

Plants were brought in as one double stemmed plant per rockwool (as expected

before the first run), and with six plants per section. This differs from the

previous run, but the statistics hold despite this inconsistency, since the number of

plants in a treatment is part of the error term for the statistical model. The plants

were brought in on December 31st in sub-zero weather, and some plants (primary

leaves) were severely damaged by the cold (Control plants, 19:1 Med and 10:1

Med sections were the most affected). This freezing damage could explain some

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of the reduction observed between these three treatments from the first and second

runs.

The heating system failed two days in February (explaining the extreme low

temperatures felt in the greenhouse), leaving extremely low temperatures in the

greenhouse. This cold impacted all plants in the greenhouse similarly, thus did

not have an impact on the statistics. The cooling system also broke down during

6 hours on the hottest day in February, as well as on the hottest days in March and

April (explaining the extreme high temperatures seen in the greenhouse). Both

the cold and the heat caused a slight amount of flower abortion, as seen during the

first run (albeit much less). The fan nearest the 10:1 low section would not close

when it was off, so cold air was allowed in. The same problem was observed with

the fan nearest the HPS section, during the coldest day in February. Both were

fixed within a few days. No impact was observed, except for the HPS section,

where 2 plants were observed to have been slightly damaged by the cold air.

They had stunted growth and less fruit than the others in the section. These 2

plants were randomly chosen to be harvested for the 70 day harvest, which

occurred less than a week after the symptoms were noticed, reducing the HPS

production values relative to the other 70 day harvests.

Similar problems from the first run were encountered, such as the incidence of the

sun (which is much lower during the winter). This might have a small effect on

production. The shade cloth has been partially removed in order to compensate

(no shade cloth above the plants). This could have resulted in the increased

results from 19:1 High, and 100% Red, since the added biomass growth due to

more red light permitted them to grow taller quicker. The lamps continued to

have voltage drops as experienced in the first run. This was corrected every two

weeks, by readjusting the voltage to the correct values for each lamp. The impact

was not statistically significant.

The 5:1 Med bulbs were unplugged three times for a total of 4 days. 19:1 Low

was broken for 15 days and was replaced. No impact on the statistics was shown.

The HPS lamp in the 50% section would shut off after a few hours during the first

two weeks of the run and was changed.

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Water lines would disconnect for some plants, due to the lines being older and

stiffer. Reconnecting the line and adding water fixed the problem, since each

plant had a water line, water could move between plants through the larger

rockwool cubes (no plant was ever completely dry).

One lamp in the 19:1 MED section fell due to a faulty wire holding it up. It

slightly damaged a plant on its way down and completely severed another. This

plant had to be harvested and was added in the 70 day report, making for four

plants in the 70 day harvest and two plants in the 120 day harvest.

6.8. Observations for future research

A number of observations were made during the experiment which would require

further research to understand what the driving influence of the results was.

A large number of clusters were seen turning into secondary stems or leaves. This

phenomenon is typically associated with a lack of light (Savoura unpublished

data), undermining tissue dissociation. This lack of light intensity should only

happen on aborted or small fruit clusters. However, it was noticed under LED

treatments with full fruit clusters, where the fruit continued to grow normally.

These results (Figure 20 below) do not agree with what Savoura expected, and

since it was only noticed on LED treatments, further research would need to be

performed to explain the phenomenon.

The lower and older leaves under the LED treatments had numerous spotting and

were more brittle, called intumescences (Rud 2009). These spots are typical of

Solanaceous crops, are cultivar dependent, and are characterized by individual

epidermal cells swelling and bursting. The process is non-reversible, and has

shown to appear on tomato plants from the Maxifort variety used as the base for

the plants (Rud 2009), but not on the Trust variety. Minor occurrences of

intumescences do not impact production or growth, but more severe cases lead to

necrosis of the leaves and can dampen the overall production by up to 50% in

extreme cases (Rud 2009).

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Figure 20: Two examples of viable clusters turning into secondary stems.

(harvested during the second run)

Intumescences were observed in all LED treatments, but not in the HPS or control

treatments. Higher intensity treatments were more severely impacted, but the

intumescences were relatively mild compared to what is shown in the literature

(Rud 2009). The worst cases are depicted in Figure 21, and should not have

impacted fruit production too heavily, since they were only observed on older

leaves and did not take up over 50% of the leaves. From these results, these spots

could be an interaction between the nutrient solution and the LED lighting. An

improved nutrient mix might have to be developed in order to compensate for this

interaction. Intumescences have been shown to occur when the moisture uptake

by the plant is higher than what the plant can evacuate (due to low vapor pressure

deficit or over watering of the leaves) (Rud 2009), which could suggest that the

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LED light promotes water uptake by the rooting system, since all plants were

submitted to the same watering conditions and only the LED treatments were

affected.

Figure 21: Leaf Burn. Observed on older leaves under the LED light fixtures

The regression curves resulted in strong fits for linear curves when applied to the

factorial models. From these results, higher ratios (i.e. 1:1 red to blue) and higher

intensities should be tested to determine the potential maximums.

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Powdery mildew was not observed during the second run but was observed in the

first run. To reduce contamination issues, a delay of over a month was allowed

between experimental runs to reduce the risk of disease carry-over. During the

first run, the 5:1 Med ratio had the most severe case of powdery mildew

potentially due to an over-exertion of the plant’s fruiting capabilities. Powdery

Mildew is rare in tomato strains which are resistant (such as the one used for this

experiment), but can occur due to exposure (Jones et al. 2001). Since no exposure

was expected, further research could determine if the overexertion theory is

correct.

Figure 22: Powdery Mildew. 5:1 Med section, during the first run.

For future research, it would be interesting to test a factorial comparison between

the 50%:50% LED:HPS and the 5:1 ratio to obtain regression curves across

increasing intensities of light.

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Future research would need to include quality (color, size, shape, taste, Brix, etc.)

with the yield data to provide a more complete evaluation of marketable fruit

quality beyond size that was reported in this document. An investigation into fruit

quality should be planned (the concentration of carotenoids in the fruit dependent

on the treatments), as well as a nutrient analysis of leaf samples, in order to

understand if a specific nutrient was also absorbed more readily in the plants

under LEDs.

A major issue from this experiment that needs to be addressed by all players

working with LED lights for horticultural purposes is the uniformity and accuracy

of light measurement techniques, equipment, and calculations for LEDs. Methods

and equipment used to measure other lighting sources (solar, HPS, incandescent,

etc.) are not consistent when measuring LED light arrays.

It can be theorized that a better sensor for LED arrays could be achieved by

placing multiple spectroradiometer type sensors in a bulb formation, in order to

get an accurate measurement of light coming from every direction, instead of a

bulb which refracts all light incoming into a single point.

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7. CONCLUSIONS

The most obvious conclusions that can be made from this experiment are that the

top five fruit producing light treatments were 5:1 High, 5:1 Med, 19:1 High, Red

Top and 50% LED:50%HPS. However, statistically no difference was measured

between any of the treatments, with the exception of the control which was

statistically different from all of the high producing treatments. The high

irradiance level was the top producer for each ratio, with the largest fresh and dry

vegetative biomass occurring within the 19:1 High LED. The top three treatments

for the largest mass of total fruit were 5:1 High, 5:1 Med, and 19:1 High. The top

three treatments of total number of fruit were 5:1 High, 5:1 Med, and 5:1 Low.

The top three treatments for marketable number of fruit were 50%:50%, Red Top

and 5:1 Med. The top three for total marketable fruit mass was 50%:50%, 5:1

High, and 19:1 High. The differences between these top three treatments (and

sometimes the top 6) were not statistically different. From this data, the top LED

treatment recommended is the 5:1 High, but the 50%:50% surpassed the 5:1 High

treatment for marketable fruit, meaning it can also be considered. From the

regression analysis of the factorial experiment, it can be reported that higher

levels of light increased production, while increased levels of red light (relative to

blue) resulted in less fruit. The regression also reported that increased levels of

light resulted in increased biomass, and increased red light (relative to blue)

resulted in more biomass. The treatment recommended for a newly built

greenhouse is the 5:1 High LED treatment (consistently found to be ahead in total

yields). For a pre-existing greenhouse however, the recommended treatment

should be the 50%:50% LED:HPS. This is due to this treatment not performing

significantly different than the 5:1 treatments, but not requiring complete removal

of the HPS lights, lowering renovation costs while still increasing overall

production significantly. The conclusions reached in this report were not based

on any financial consideration but only on the results from the test and time of

installation of the different systems within the greenhouse. Overall, it was shown

that the top LED treatments (5:1 High, 5:1 Med and 19:1 High) as well as the

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50%:50% treatment consistently outperformed the HPS treatment, and thus these

treatments can be considered an improvement over traditional HPS lighting for

greenhouses.

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Apogee Instruments Inc. 2012. Apogee Quantum Sensor Technical Information. www.apogeeinstruments.co.uk/apogee-instruments-quantum-sensor-technical-information/

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APPENDIX A: Formatted Data

Figure A1: Total marketable fruit for the second run for the tomato plants.

Figure A2: Total marketable fruit for the first run for the tomato plants.

0

10

20

30

40

50

60

70

80

90

5:1

LO

W

5:1

MED

5:1

HIG

H

10

:1 L

OW

10

:1 M

ED

10

:1 H

IGH

19

:1 L

OW

19

:1 M

ED

19

:1 H

IGH

LED

RED

TO

P

RED

BO

T

50

/50

HP

S

CO

NTR

OL

Seco

nd

ru

n t

ota

l nu

mb

er

of

mar

keta

ble

fru

it

70 day

120 day

Total

0

5

10

15

20

25

30

35

40

45

50

5:1

LO

W

5:1

MED

5:1

HIG

H

10

:1 L

OW

10

:1 M

ED

10

:1 H

IGH

19

:1 L

OW

19

:1 M

ED

19

:1 H

IGH

LED

RED

TO

P

RED

BO

T

50

/50

HP

S

CO

NTR

OL

Firs

t ru

n t

ota

l nu

mb

er

of

mar

keta

ble

fru

it

70 day

120 day

Total

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Figure A3: Total mass of marketable fruit for the second run for the tomato plants.

Figure A4: Total mass of marketable fruit for the first run for the tomato plants.

0

2000

4000

6000

8000

10000

12000

14000

Seco

nd

ru

n t

ota

l mas

s o

f m

arke

tab

le f

ruit

(g)

70 day

120 day

Total

0

1000

2000

3000

4000

5000

6000

Firs

t ru

n t

ota

l mas

s o

f m

arke

tab

le f

ruit

(g)

70 day

120 day

Total

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Figure A5: Total fruit count during the second run

Figure A6: Total fruit count during the first run. * The HPS conjecture is overestimated; since it keeps into account the 20% increase during the 70 day run while for the 120 day run it would be closer to 5% increase.

0

20

40

60

80

100

120

140

160

180

200

Seco

nd

ru

n t

ota

l fru

it c

ou

nt

70 day

120 day

Total

0

20

40

60

80

100

120

140

160

180

200

5:1

LO

W

5:1

MED

5:1

HIG

H

10

:1 L

OW

10

:1 M

ED

10

:1 H

IGH

19

:1 L

OW

19

:1 M

ED

19

:1 H

IGH

LED

RED

TO

P

RED

BO

T

50

/50

HP

S

CO

NTR

OL

Fruit Count

Firs

t ru

n t

ota

l fru

it c

ou

nt

70 day

120 day

Total

HPS Conjecture

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Figure A7: Total fruit mass during the second run

Figure A8: Total fruit mass during the first run

* The HPS conjecture is overestimated; since it keeps into account the 20% increase during the 70 day run while for the 120 day run it would be closer to 5% increase.

0

2000

4000

6000

8000

10000

12000

14000

16000

18000

20000

Seco

nd

ru

n t

ota

l fru

it m

ass

(g)

70 day

120 day

Total

0

2000

4000

6000

8000

10000

12000

5:1

LO

W

5:1

MED

5:1

HIG

H

10

:1 L

OW

10

:1 M

ED

10

:1 H

IGH

19

:1 L

OW

19

:1 M

ED

19

:1 H

IGH

LED

RED

TO

P

RED

BO

T

50

/50

HP

S

CO

NTR

OL

Fruit Weight

Firs

t ru

n t

ota

l fru

it m

ass

(g)

70 day

120 day

Total

HPS Conjecture

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Figure A9: Total plant fresh biomass (excluding fruit) for the second run

Figure A10: Total plant fresh biomass (excluding fruit) for the first run

0

5000

10000

15000

20000

25000

30000

Seco

nd

ru

n t

ota

l fre

sh b

iom

ass

(g)

70 day

120 day

Total

0

2000

4000

6000

8000

10000

12000

14000

16000

18000

Firs

t ru

n t

ott

al f

resh

bio

mas

s (g

)

70 day

120 day

Total

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Figure A11: Total plant dried biomass (excluding fruit) for the second run

Figure A12: Total plant dried biomass (excluding fruit) for the first run.

APPENDIX B: Statistical Data (Box Plots)

0

200

400

600

800

1000

1200

1400

1600

1800

Seco

nd

ru

n t

ota

l dry

bio

mas

s (g

)

70 day

120 day

Total

0

200

400

600

800

1000

1200

1400

5:1

LO

W

5:1

MED

5:1

HIG

H

10

:1 L

OW

10

:1 M

ED

10

:1 H

IGH

19

:1 L

OW

19

:1 M

ED

19

:1 H

IGH

LED

RED

TO

P

RED

BO

T

50

/50

HP

S

CO

NTR

OL

Dry Biomass

Firs

t ru

n t

ota

l dry

bio

mas

s (g

)

70 day

120 day

Total

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For the following Figures, the mean is represented by a diamond shape, the median as a horizontal line, the lower quartile and the upper quartile as the ends of the boxes, the whiskers as the lowest and highest value observed, and circles for outliers.

Figure B1: Total number of marketable fruit per plant for the 120 day harvests

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Figure B2: Total marketable fruit mass per plant, for the 120 day harvests

Figure B3: Total number of fruit per plant, for the 120 day harvests

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Figure B4: Total mass of fruit per plant, for the 120 day harvests

Figure B5: Total fresh biomass per plant for the 120 day harvests

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Figure B6: Total dry biomass per plant for the 120 day harvests

Figure B7: Ratio of dry to fresh biomass production

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Figure B8: Ratio of fruit mass to fresh biomass for the 120 day data

Figure B9: Ratio of marketable fruit mass to fresh biomass for the 120 day data

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Figure B10: Total number of red fruit per plant, for the 120 day experiment

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APPENDIX C: Raw Data Table C1: Total fruit data (green and red fruit for the 2 replications (4 plants harvested after 70 and 120 days for the first one, and 3 plants harvested at 70 and 120 days for the second).

Fruit Count

Total Fruit Mass

Average Mass

Fruit Count

Total Fruit Mass

Average Mass

Fruit Count

Total Fruit Mass

Average Mass

Run 1 5:1 LOW 5:1 MED 5:1 HIGH 70 day 58 1549.2 26.71 92 3355.6 36.47 84 2541.8 30.26 120 day 115 7998.8 69.55 100 7896.5 78.97 107 7866.9 73.52

Total 173 9548 55.19 192 11252.1 58.6 191 10408.7 54.5 Run 2 70 day 46 1327.6 28.86 33 1016.1 30.79 55 1909.3 34.71 120 day 122 11727 96.12 133 13557.3 101.93 139 14908.5 107.26

Total 168 13054.6 77.71 166 14573.4 87.79 194 16817.8 86.69 Run1 10:1 LOW 10:1 MED 10:1 HIGH 70 day 13 342 26.31 46 2682.6 58.32 54 2625.3 48.62 120 day 75 4277.6 57.03 60 4131.1 68.85 107 7033.3 65.73

Total 88 4619.6 52.5 106 6813.7 64.28 161 9658.6 59.99 Run 2 70 day 34 849.7 24.99 52 1927.2 37.06 44 1612.2 36.64 120 day 84 7456.2 88.76 97 9696.1 99.96 99 10293.2 103.97

Total 118 8305.9 70.39 149 11623.3 78.01 143 11905.4

83.25

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Run 1 19:1 LOW 19:1 MED 19:1 HIGH 70 day 12 526.2 43.85 28 1055.8 37.71 54 2026.2 37.52 120 day 41 2470.1 60.25 71 4286.6 60.37 64 4560.5 71.26

Total 53 2996.3 56.53 99 5342.4 53.96 118 6586.7 55.82 Run 2 70 day 39 1138 29.18 21 582.8 27.75 69 2483.8 36

16 489.5 30.59 120 day 105 9432.4 89.83 85 8895.7 104.66 128 14885 116.29

Total 105 9432.4 89.83 101 9385.2 92.92 128 14885 116.29 Run 1 LED RED TOP RED BOT 70 day 46 1465.8 31.87 34 1465.7 43.11 42 1427.1 33.98 120 day 114 7225.5 63.38 78 6605.9 84.69 74 4600.8 62.17

Total 160 8691.3 54.32 112 8071.6 72.07 116 6027.9 51.96 Run 2 70 day 56 1967 35.13 26 1072.1 41.23 32 759 23.72 120 day 94 10493.8 111.64 130 14170.4 109.00 120 13941.8 116.18

Total 150 12460.8 83.07 156 15242.5 97.71 152 14700.8 96.72 50/50 HPS CONTROL

70 day 42 1608.3 38.29 16 378.3 23.64 0 0 --- 120 day 64 5452.7 85.2 37 3018 81.57 13 1190.8 91.6

Total 106 7061 66.61 53 3396.3 64.08 13 1190.8 91.6 Run 2 70 day 54 1796.6 33.27 4 88 22 0 0 --- 120 day 109 14302.5 131.22 92 9069.3 98.58 10 624.5 62.45

Total 163 16099.1 98.77 96 9157.3 95.39 10 624.5 62.45

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A fourth plant had to be weighed in the 19:1 MED section, since a lamp fell and killed it. The values in red are the sum of all four plants, while the values underneath were found using the best, the worst and the average of the two middle plants. (The 120 data for 19:1 Med therefore only includes 2 plants) Table C2: Red fruit harvested data during the 120 day experiment (4 plants harvested at 70 days, 4 at 120 days). Red fruit were harvested from all plants when the first red pigmentation was noticed on the green fruit. No red fruit was harvested at the 70 day mark during the run 2.

Run 1 5:1 LOW

5:1 MED 5:1 HIGH

70 day 3 9 2 120 day 78 72 73

Total 81 81 75 Run 2

120 day 13 20 39

Run 1 10:1 LOW

10:1 MED 10:1 HIGH

70 day 0 7 6 120 day 36 45 72

Total 36 52 78 Run 2

120 day 2 16 18

Run1 19:1 LOW

19:1 MED 19:1 HIGH

70 day 5 0 1 120 day 17 44 46

Total 22 44 47 Run 2

120 day 10 15 21

Run 1 LED RED TOP RED BOT

70 day 7 5 4 120 day 73 59 35

Total 80 64 39 Run 2

120 day 10 28 7

Run 1 50/50 HPS CONTROL 70 day 2 0 0 120 day 48 19 7

Total 50 19 7 Run 2

120 day 23 2 0

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Table C3: Greater than 90 g fruit data during the first (4 plants harvested at 70 days and 4 at 120 days) and the second (3 plants harvested at 70 days) replications. The total number of fruit (count) from each treatment and mass of these fruit is recorded.

Run 1 5:1 LOW

5:1 MED 5:1 HIGH

Count 37 39 33 Mass 4057.4 4399.2 4049.4 Run 2 Count 55 70 70 Mass 8431.5 10745.4 12167.4

Run 1 10:1 LOW

10:1 MED 10:1 HIGH

Count 17 19 21 Mass 1900.2 2105.2 2315.7 Run 2 Count 36 54 63 Mass 5246.7 8138.6 8576

Run 1 19:1 LOW

19:1 MED 19:1 HIGH

Count 13 13 27 Mass 1404.7 1349.2 2905.1 Run 2 Count 47 44 81 Mass 7127.4 7231.1 13162.8

Run 1 LED RED TOP RED BOT

Count 26 32 20 Mass 2803.2 3868.4 2433.9 Run 2 Count 59 79 62 Mass 9539 11535.6 11856.5 Run 1 50/50 HPS CONTROL Count 37 14 5 Mass 4166.6 1673.5 735.4 Run 2 Count 77 48 10 Mass 12794.8 7442.4 624.5

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Table C4: Biomass data (fresh and dry) during the first (4 plants harvested at 70 days and 4 at 120 days) and the second (3 plants harvested at 70 days) replications.

Fresh Mass

Average Plant

Dry Mass

Average Plant

Fresh Mass

Average Plant

Dry Mass

Average Plant

Fresh Mass

Average Plant

Dry Mass

Average Plant

Run 1 5:1 LOW 5:1

MED 5:1 HIGH

70 day 3857.2 964.3 279.4 69.9 4961.4 1240.3 354.7 88.7 5060.3 1265.1 402.3 100.6 120 day 7001.1 1750.3 526.1 131.5 7838.7 1959.7 569.3 142.3 10016.9 2504.2 713 178.3

Total 10858.3 1357.3 805.5 100.7 12800.1 1600 924 115.5 15077.2 1884.6 1115.3 139.4 Run 2 70 day 6129.3 2043.1 423.1 141 5501.6 1833.867 358.9 119.6 6414.5 2138.167 414.1 138 120 day 8983.2 2994.4 755.7 251.9 14587.8 4862.6 1141.3 380.4333 11827.3 3942.433 1043.8 347.9333

Total 15112.5 2518.75 1178.8 196.47 20089.4 3348.23 1500.2 250.03 18241.8 3040.30 1457.9 242.98

Run 1 10:1 LOW 10:1

MED 10:1 HIGH

70 day 4212.2 1053.1 285.2 71.3 5142.9 1285.7 350.8 87.7 4585 1146.3 371.8 93 120 day 10000 2500 709.1 177.3 8794.1 2198.5 583.8 146 10677.5 2669.4 825.9 206.5

Total 14212.2 1776.5 994.3 124.3 13937 1742.1 934.6 116.8 12832.8 1604.1 992.9 124.1 Run 2 70 day 5341.7 1780.567 407 135.7 6365.9 2121.967 348 116 6152.9 2050.967 368.9 123 120 day 14826.3 4942.1 1105.9 368.6333 14357.9 4785.967 1221.6 407.2 11208.6 3736.2 829.8 276.6

Total 20168 3361.33 1512.9 252.15 20723.8 3453.97 1569.6 261.60 17361.5 2893.58 1198.7 199.78

Run 1 19:1 LOW 19:1

MED 19:1 HIGH

70 day 4117.2 1029.3 268.2 67.1 4163 1040.8 282.8 70.7 5995.5 1498.9 449.9 112.5 120 day 8371.2 2092.8 580.9 145.2 11868.2 2967.1 850.4 212.6 10939.4 2734.9 836.3 209.1

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Total 12488.4 1561.1 849.1 106.1 16031.2 2003.9 1133.2 141.7 16934.9 2116.9 1286.2 160.8 Run 2 70 day 5653.1 1884.367 355.7 118.6 7653.8 1913.45 544.3 136.1 6404.7 2134.9 449.1 149.7

4602.5 1534.167 334.9 111.6 120 day 15782.4 5260.8 1284.2 428.0667 10759.7 3586.567 906.3 302.1 15553.3 5184.433 1092 364

Total 21435.5 3572.58 1639.9 273.32 18413.5 3068.92 1450.6 241.77 21958 3659.67 1541.1 256.85

Run 1 100% LED RED

TOP RED BOT

70 day 5216.1 1304 365.8 91.5 4783.2 1195.8 302.9 75.7 3915.7 978.9 309.4 77.4 120 day 10395.8 2599 763.9 191 9753.3 2438.3 671.9 168 9951.6 2487.9 640.5 160.1

Total 15611.9 1951.5 1129.7 141.2 14536.5 1817.1 974.8 121.9 13867.3 1733.4 949.9 118.7 Run 2 70 day 8058 2686 546 182 4477.8 1492.6 295 98.3 5827.7 1942.567 426.1 142 120 day 13233 4411 1088.6 362.8667 12250.5 4083.5 961.4 320.4667 18947.2 6315.733 1108.9 369.6333

Total 21291 3548.50 1634.6 272.43 16728.3 2788.05 1256.4 209.40 24774.9 4129.15 1535 255.83 Run 1 50/50 HPS CONT 70 day 4333 1083.3 311.6 77.9 3978.3 994.6 251.9 63 1590.1 397.5 89.8 22.5 120 day 9577.1 2394.3 667.7 166.9 6783 1695.8 441.8 110.5 3879.7 969.9 231.3 57.8

Total 13910.1 1738.8 979.3 122.4 10761.3 1345.2 693.7 86.7 5469.8 683.7 321.1 40.1 Run 2 70 day 5688.9 1896.3 365 121.7 4118.4 1372.8 306.5 102.2 1245.9 415.3 68.9 23 120 day 15649 5216.333 1037 345.6667 12774.6 4258.2 1414.1 471.3667 8118 2706 604.5 201.5

Total 21337.9 3556.32 1402 233.67 16893 2815.50 1720.6 286.77 9363.9 1560.65 673.4 112.23

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A fourth plant had to be weighed in the 19:1 MED section at the 70 day mark of the second run, since a lamp fell and killed it. The values in red are the sum of all four plants, while the values underneath were found using the best, the worst and the average of the two middle plants.

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Table C5: Fruit and flower counts for both replications. The total counts are provided

Date 5:1 LOW

5:1 MED

5:1 HIGH

Run 1 Flower Fruit Flower Fruit Flower Fruit 2-Aug 27 10 25 36 26 15 9-Aug 38 14 61 42 62 29

12-Aug 58 25 49 47 56 44

16-Aug 46 39 38 68 30 62

25-Aug 44 76 34 102 36 99

1-Sep 24 115 14 148 32 148 28-Sep 13 74 22 72 19 59 Run 2 8-Feb 103 20 82 10 111 28 28-Feb 80 78 86 92 119 130 27-Apr 52 122 138 133 126 139

10:1 LOW

10:1 MED

10:1 HIGH

Run 1 Flower Fruit Flower Fruit Flower Fruit 2-Aug 43 2 40 29 23 12 9-Aug 41 4 46 32 66 25

12-Aug 38 5 37 37 38 34

16-Aug 19 8 24 42 34 54

25-Aug 34 44 15 69 30 101

1-Sep 31 44 10 95 12 148 28-Sep 19 44 5 29 30 64 Run 2 8-Feb 77 5 106 19 111 32 28-Feb 66 56 84 81 98 90 27-Apr 235 84 290 97 139 99

19:1 LOW

19:1 MED

19:1 HIGH

Run 1 Flower Fruit Flower Fruit Flower Fruit 2-Aug 36 3 34 23 32 16 9-Aug 28 9 41 29 49 17

12-Aug 23 10 26 31 36 34

16- 8 12 24 36 31 42

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Aug 25-Aug 18 32 27 47 26 69

1-Sep 16 34 9 80 16 92 28-Sep 10 27 29 25 15 43 Run 2 8-Feb 81 8 47 8 93 28 28-Feb 73 72 68 38 88 104 27-Apr 165 105 189 85 167 128

LED

RED TOP

RED BOT

Run 1 Flower Fruit Flower Fruit Flower Fruit 2-Aug 72 17 29 11 23 13 9-Aug 33 31 23 18 29 11

12-Aug 27 34 25 18 26 15

16-Aug 20 46 21 17 23 40

25-Aug 32 95 30 66 19 56

1-Sep 33 114 20 82 16 72 28-Sep 30 65 12 70 23 40 Run 2 8-Feb 101 13 80 31 81 4 28-Feb 101 90 63 91 111 64 27-Apr 331 94 203 130 231 120

50/50 HPS

CONTROL

Run 1 Flower Fruit Flower Fruit Flower Fruit 2-Aug 22 12 2 0 2 0 9-Aug 27 16 27 0 13 0

12-Aug 34 20 35 3 13 3

16-Aug 27 27 17 4 6 2

25-Aug 42 64 34 30 19 10

1-Sep 24 85 38 33 12 9 28-Sep 24 40 27 49 18 11 Run 2 8-Feb 90 25 38 2 0 0 28-Feb 87 112 57 15 4 0 27-Apr 266 109 138 92 86 10

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APPENDIX D: Weather and Lighting Data

Table D1: Temperature data during the second run

5:1 LOW 5:1 MED 5:1 HIGH Temp SD MAX MIN Temp SD MAX MIN Temp SD MAX MIN January 16.8 2.7 24.4 10.4 16.5 2.3 23.4 5.3 16.2 2.4 23.6 10.5 February 15.8 4.1 28.6 1.7 16 4.2 28.2 2 15.1 3.7 27.1 1.8 March 17.4 3.7 34.5 9.7 17.3 3.5 29.7 9.9 17.3 3.4 28.9 8.7 April 17.8 4 34.3 10.6 17.4 3.6 32.6 10.9 18.2 3.9 39.2 10.9 10:1 LOW 10:1 MED 10:1 HIGH Temp SD MAX MIN Temp SD MAX MIN Temp SD MAX MIN January 16.2 2.4 23.6 10.5 16.6 2.5 23.6 8.7 16 2.3 22.5 5.9 February 15.1 3.7 27.1 1.8 15.7 4.2 28.9 1.8 15.7 4 26.9 2.5 March 17.9 3.9 33.5 9.2 17.6 3.6 29.5 10 17.2 3.7 30.9 10.1 April 18.4 4.2 40.2 9.1 18.5 4 39.3 10.7 17.4 3.6 32.5 10.8 19:1 LOW 19:1 MED 19:1 HIGH Temp SD MAX MIN Temp SD MAX MIN Temp SD MAX MIN January 16.7 2.6 24.3 10.4 16.9 2.6 25.3 10.6 16.1 2.3 22.1 5 February 16.1 4.2 28.8 1.6 17.2 2.3 20.4 2.5 15.8 4 27 2.2 March 17.6 3.9 34.5 9.6 18.2 4 31.2 10.3 17 3.5 29.2 9.5 April 18.3 4.3 36 10.5 18.6 4.1 38.4 10.7 17.3 3.6 32.6 10.6

LED RED TOP RED BOT Temp SD MAX MIN Temp SD MAX MIN Temp SD MAX MIN January 16.4 2.4 23.6 10.1 16.3 2.3 22.8 5.8 16.7 2.6 22.9 10.3 February 16.6 4 29.3 2.4 16.2 4.1 28.1 2.2 15.6 3.8 27.2 2 March 17.7 3.9 31.1 9.6 17.4 3.7 30.7 9.9 16.8 3.4 29.3 9.3 April 17.4 4.3 39.5 10.8 17.7 3.9 33.9 11 16.7 3.3 31.6 10.1 50/50 HPS CONTROL Temp SD MAX MIN Temp SD MAX MIN Temp SD MAX MIN January 16.2 2.4 23.2 5 16.7 2.8 26.3 7.3 16.7 2.5 24.9 11 February 15.9 4 28.1 2 15.9 4.3 31.5 0.9 15.9 3.9 29.4 2 March 17.2 3.6 31.7 9.4 16.9 3.6 31.1 2.5 16.8 3.1 29.2 9.7 April 17.6 3.8 33.7 10.6 16.3 3.2 31.6 10.4 17.3 3.4 35.6 10

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Table D2: Temperature data during the first run 5:1 LOW 5:1 MED 5:1 HIGH

Temperature St Dev Max Min Temperature St

Dev Max Min Temperature St Dev Max Min

21.4 2.1 33.6 11.2 23.1 2.0 33.1 16.1 21.3 2.1 32.9 11.1 10:1 LOW 10:1 MED 10:1 HIGH

Temperature St Dev Max Min Temperature St

Dev Max Min Temperature St Dev Max Min

22.1 1.8 33.3 9.9 23.2 2.2 32.6 16.4 20.7 2.8 32.2 10.5 19:1 LOW 19:1 MED 19:1 HIGH

Temperature St Dev Max Min Temperature St

Dev Max Min Temperature St Dev Max Min

21.9 1.7 35.2 10.8 23.0 2.1 31.7 16.3 21.4 2.1 35.2 11.1 LED RED TOP RED BOT

Temperature St Dev Max Min Temperature St

Dev Max Min Temperature St Dev Max Min

21.2 2.2 31.2 10.5 20.9 2.5 35.8 10.4 21.5 2.6 38.7 10.8 50/50 HPS CONTROL

Temperature St Dev Max Min Temperature St

Dev Max Min Temperature St Dev Max Min

21.2 2.2 33.5 11.1 21.3 3.0 38.5 9.9 21.0 3.1 36.9 10.7

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Table D3: End of second run light map data (May 1st 2012)

Zone Spectroradiometer Li-COR Daily Light Integral (mol day-1)

% of Artificial

Light Average Vertical

Average 45º angle Average Average

5:1 LOW 98.2 152.9 125.6 118.1 8.5 37.3 5:1 MED 119.1 154.2 136.7 152.4 9.8 40.6 5:1 HIGH 130.3 154.2 142.3 190.9 11.5 44.5 10:1 LOW 98.6 126.6 112.6 113.7 9.1 38.8 10:1 MED 113.2 148.7 130.9 134.3 9.9 40.9 10:1 HIGH 125.2 151.4 138.3 160.7 13.1 47.9 19:1 LOW 98.4 143.9 121.2 112.8 9 38.7 19:1 MED 114.4 141.7 128.1 139.7 10.4 42.2 19:1 HIGH 125.3 144 134.7 177.6 12 45.7

LED 131 163 147 206.6 12.3 46.2 0.5 114.7 142 128.3 110.9 9.1 38.8

Red TOP 119.3 150.2 134.8 147.6 11.5 44.6 Red BOT 104 137.1 120.6 115.1 7.9 35.5 Control 0.4 0.1 0.3 0.6 0 0.1

HPS Light Maximum Irradiance Middle (1 m) 38 38 37.3 4.4 23.4 Height (1.5 m) 96 96 127.8 9.2 39.2 Top (30 cm from lamp) 162.6 162.6 447.7 21.9 60.5

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Table D4: Initial light map data (August 11th 2011), with correction from updated calibration (next page)

LED Light Maps Average Vertical

Average 45º

angle Horizontal Average

Daily Light Integral (mol

day-1)

% of Artificial

Light

5:1 LOW 51.625 71.875 13.75 61.75 5.34 27.2 5:1 MED 61 87 14.25 74 6.39 30.9 5:1 HIGH 70.125 106.25 29.875 88.125 7.61 34.7 10:1 LOW 73.875 97.25 20.625 85.5 7.39 34.1 10:1 MED 78.875 96.75 33.875 87.875 7.59 34.7 10:1 HIGH 83.875 105.25 23 94.625 8.18 36.4 19:1 LOW 67.375 91.625 19.5 79.5 6.87 32.4 19:1 MED 71.125 93.375 25.75 82.25 7.11 33.2 19:1 HIGH 91.25 104.375 22.375 97.875 8.46 37.2

LED 69.75 97.25 116.75 83.5 7.21 33.5 RED TOP 78 102.125 73.75 90 7.78 35.2 RED BOT 74 88.25 38 81.125 7.01 32.9

50/50 58.5 79.375 39 69 5.96 29.4 HPS Light Starting Irradiance Maximum Irradiance

Floor 13.5 10.875 Middle (1 m) 42.625 37.5

Plant height(1.5 m) 30 75 6.48 31.2 Top (30 cm from

lamp) 260 175

Light map was for the LED bulbs in a vertical and 45 degree angle at a distance of 30 cm from the bulb in the center of the chamber at three locations for each vertical and angled measurement (full darkness, with other bulbs on). The HPS was measured at three heights (top of rock wool cube, middle of the chamber (approximate height of the seedlings), and top of the chamber (30 cm from the bulb in a vertical orientation). All measurements were taken with a spectroradiometer, corrected from calibration on Dec 15, 2011.

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Table D5: Initial summary of light map data (August 11th 2011), without calibration correction

5:1 LOW 5:1 MED 5:1 HIGH PAR VALUES (μmol m-2 s-1) PAR VALUES (μmol m-2 s-1) PAR VALUES (μmol m-2 s-1) AVERAGE 49.4 AVERAGE 59.2 AVERAGE 70.5 10:1 LOW 10:1 MED 10:1 HIGH PAR VALUES (μmol m-2 s-1) PAR VALUES (μmol m-2 s-1) PAR VALUES (μmol m-2 s-1) AVERAGE 68.4 AVERAGE 70.2 AVERAGE 75.7 19:1 LOW 19:1 MED 19:1 HIGH PAR VALUES (μmol m-2 s-1) PAR VALUES (μmol m-2 s-1) PAR VALUES (μmol m-2 s-1) AVERAGE 63.6 AVERAGE 65.8 AVERAGE 78.3 LED RED TOP RED BOT PAR VALUES (μmol m-2 s-1) PAR VALUES (μmol m-2 s-1) PAR VALUES (μmol m-2 s-1) AVERAGE 66.9 AVERAGE 72 AVERAGE 65.9 50/50 HPS CONTROL PAR VALUES (μmol m-2 s-1) PAR VALUES (μmol m-2 s-1) PAR VALUES (μmol m-2 s-1)

AVERAGE 55.2+(24 from HPS) AVERAGE 24 AVERAGE 0

Average after change 60

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Table D6: GE Spectroradiometer data. End of run 1 and start of run 2 light map data (January 5th 2012) Zone GE Spectroradiometer

Average Vertical

Average 45º angle

Average

5:1 LOW 692.5 302.0 497.3 5:1 MED 654.3 861.4 757.9 5:1 HIGH 963.6 1056.1 1009.9 10:1 LOW 673.9 402.9 538.4 10:1 MED 692.3 663.4 677.9 10:1 HIGH 966.2 743.5 854.9 19:1 LOW 614.3 546.0 580.1 19:1 MED 757.1 694.0 725.5 19:1 HIGH 846.9 836.1 841.5

LED 829.0 583.0 706.0 50% 912.5 509.2 710.9

Red TOP 905.0 465.2 685.1 Red BOT 738.3 541.8 640.0 Control 1.0 1.0

HPS Light Middle (1 m) 918.6 918.6

Height (1.5 m) 1045.0 1045.0 Top (30 cm from

lamp) 2106.9 2106.9

Light map was for the LED bulbs in a vertical and 45 degree angle at a distance of 30 cm from the bulb in 9 locations per section (full darkness, with other bulbs on). The HPS was measured at three heights (of the chamber (approximate height of the seedlings), top of the chamber and 30 cm from the bulb (in a vertical orientation). All measurements were taken with a spectroradiometer.


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