Photobiology in protected horticulture

The introduction of high power LED lighting systems for horticulture has stimulated substantial interest from both the research community and the protected horticulture industry. LED lighting systems have the potential to reduce electrical energy consumption compared to conventional high pressure sodium lights and their energy efficiency continues to improve. In addition to the potential of LEDs to reduce carbon footprints and reduce running costs, LED lighting also provides considerable opportunities to exploit the wealth of photobiological knowledge to produce horticultural benefits. The narrow emission spectra of LEDs allows lighting systems to be tightly designed to stimulate specific plant photoreceptors, allowing plants to be manipulated to produce desirable characteristics. Lighting systems can be designed to maximize growth, control morphology, and optimize flavor and pigmentation. This review outlines how the light spectrum influences photosynthesis and how plant photoreceptors sense light and control growth. The review then discusses the ways in which this knowledge is being implemented in commercial horticulture to improve factors such as yield, flavor, color, plant growth, and flowering as well as pest and pathogen management and control. Research in this area is moving rapidly as the LED systems improve and increase in efficiency and as the range of novel horticultural applications expands.

[1]  Takeshi Inoue,et al.  Green light drives leaf photosynthesis more efficiently than red light in strong white light: revisiting the enigmatic question of why leaves are green. , 2009, Plant & cell physiology.

[2]  C. A. Mitchell,et al.  Supplemental lighting for greenhouse-grown tomatoes: Intracanopy led towers vs. overhead HPS lamps , 2014 .

[3]  D. C. Elliott Temperature-sensitive Responses of Red Light-dependent Betacyanin Synthesis. , 1979, Plant physiology.

[4]  M. Tevini,et al.  Some effects of enhanced UV-B irradiation on the growth and composition of plants , 1981, Planta.

[5]  Linda Chalker-Scott,et al.  Environmental Significance of Anthocyanins in Plant Stress Responses , 1999 .

[6]  S. Doveri,et al.  Aroma characterisation and UV elicitation of purple basil from different plant tissue cultures. , 2013, Food chemistry.

[7]  R. Bula,et al.  Light-emitting diodes as a radiation source for plants. , 1991, HortScience : a publication of the American Society for Horticultural Science.

[8]  R. Pierik,et al.  Canopy light cues affect emission of constitutive and methyl jasmonate-induced volatile organic compounds in Arabidopsis thaliana , 2013, The New phytologist.

[9]  Zhi-gang Xu,et al.  Effect of light-emitting diodes on growth and morphogenesis of upland cotton (Gossypium hirsutum L.) plantlets in vitro , 2010, Plant Cell, Tissue and Organ Culture (PCTOC).

[10]  O. Kooten,et al.  Plasticity of photosynthesis after the ‘red light syndrome’ in cucumber , 2016 .

[11]  J. Christie,et al.  Phototropins 1 and 2: versatile plant blue-light receptors. , 2002, Trends in plant science.

[12]  K. Folta,et al.  Contributions of green light to plant growth and development. , 2013, American journal of botany.

[13]  J. Weller,et al.  Manipulation of the Blue Light Photoreceptor Cryptochrome 2 in Tomato Affects Vegetative Development, Flowering Time, and Fruit Antioxidant Content1 , 2005, Plant Physiology.

[14]  Steven M. Reppert,et al.  Polarized Light Helps Monarch Butterflies Navigate , 2004, Current Biology.

[15]  Eva Rosenqvist,et al.  Spectral effects of supplementary lighting on the secondary metabolites in roses, chrysanthemums, and campanulas. , 2014, Journal of plant physiology.

[16]  J. K. Kim,et al.  Effects of white, blue, and red light-emitting diodes on carotenoid biosynthetic gene expression levels and carotenoid accumulation in sprouts of tartary buckwheat (Fagopyrum tataricum Gaertn.). , 2013, Journal of agricultural and food chemistry.

[17]  X. Hao,et al.  Effect of LED interlighting combined with overhead HPS light on fruit yield and quality of year-round sweet pepper in commercial greenhouse , 2016 .

[18]  C. Brown,et al.  Root-Shoot Interaction in the Greening of Wheat Seedlings Grown under Red Light , 1995, Plant physiology.

[19]  R. Pierik,et al.  Perception of low red:far-red ratio compromises both salicylic acid- and jasmonic acid-dependent pathogen defences in Arabidopsis. , 2013, The Plant journal : for cell and molecular biology.

[20]  Myung-Min Oh,et al.  Leaf Shape, Growth, and Antioxidant Phenolic Compounds of Two Lettuce Cultivars Grown under Various Combinations of Blue and Red Light-emitting Diodes , 2013 .

[21]  Jingming Zheng,et al.  Led inter-lighting in year-round greenhouse mini-cucumber production , 2012 .

[22]  Hyeon-Hye Kim,et al.  Stomatal conductance of lettuce grown under or exposed to different light qualities. , 2004, Annals of botany.

[23]  Suruchi Singh,et al.  Supplemental ultraviolet-B induced changes in essential oil composition and total phenolics of Acorus calamus L. (sweet flag). , 2009, Ecotoxicology and environmental safety.

[24]  C. Kubota,et al.  Effects of supplemental light quality on growth and phytochemicals of baby leaf lettuce , 2009 .

[25]  T. Sharkey,et al.  Efficiency of photosynthesis in continuous and pulsed light emitting diode irradiation , 1995, Photosynthesis Research.

[26]  Yun‐Hi Kim,et al.  Influences of four different light-emitting diode lights on flowering and polyphenol variations in the leaves of chrysanthemum (Chrysanthemum morifolium). , 2012, Journal of agricultural and food chemistry.

[27]  E. Runkle,et al.  An intermediate phytochrome photoequilibria from night-interruption lighting optimally promotes flowering of several long-day plants , 2016 .

[28]  M. W. Parker,et al.  A Reversible Photoreaction Controlling Seed Germination. , 1952, Proceedings of the National Academy of Sciences of the United States of America.

[29]  E. Goto,et al.  Effects of Ultraviolet Light on Growth, Essential Oil Concentration, and Total Antioxidant Capacity of Japanese Mint , 2010 .

[30]  C. Ballaré,et al.  A look into the invisible: ultraviolet-B sensitivity in an insect (Caliothrips phaseoli) revealed through a behavioural action spectrum , 2010, Proceedings of the Royal Society B: Biological Sciences.

[31]  K. Folta,et al.  Light as a Growth Regulator: Controlling Plant Biology with Narrow-bandwidth Solid-state Lighting Systems , 2008 .

[32]  T. Dougher,et al.  Differences in the Response of Wheat, Soybean and Lettuce to Reduced Blue Radiation¶ , 2001, Photochemistry and photobiology.

[33]  M. Johkan,et al.  Effects of Supplemental Lighting with Light-Emitting Diodes (LEDs) on Tomato Yield and Quality of Single-Truss Tomato Plants Grown at High Planting Density , 2012 .

[34]  A. Brazaitytė,et al.  The effect of red and blue light component on the growth and development of frigo strawberries , 2010 .

[35]  S. B. Agrawal,et al.  Supplemental UV‐B induced changes in leaf morphology, physiology and secondary metabolites of an Indian aromatic plant Cymbopogon citratus (D.C.) Staph under natural field conditions , 2010 .

[36]  J. Zeier,et al.  Light Regulation and Daytime Dependency of Inducible Plant Defenses in Arabidopsis: Phytochrome Signaling Controls Systemic Acquired Resistance Rather Than Local Defense1 , 2008, Plant Physiology.

[37]  J. Tumlinson,et al.  Plant volatiles as a defense against insect herbivores , 1999, Plant physiology.

[38]  E. Runkle,et al.  Specific functions of red, far red, and blue light in flowering and stem extension of long-day plants , 2001 .

[39]  M. Daws,et al.  Impact of red : far red ratios on germination of temperate forest herbs in relation to shade tolerance, seed mass and persistence in the soil , 2007 .

[40]  H. Kuba,et al.  The effectiveness of a green light emitting diode (LED) trap at capturing the West Indian sweet potato weevil, Euscepes postfasciatus (Fairmaire) (Coleoptera: Curculionidae) in a sweet potato field , 2004 .

[41]  Jihong Liu Clarke,et al.  Artificial light from light emitting diodes (LEDs) with a high portion of blue light results in shorter poinsettias compared to high pressure sodium (HPS) lamps , 2012 .

[42]  D. Fanourakis,et al.  Co-ordination of hydraulic and stomatal conductances across light qualities in cucumber leaves , 2011, Journal of experimental botany.

[43]  E. Goto,et al.  Effects of light quality and light period on flowering of everbearing strawberry in a closed plant production system , 2012 .

[44]  K. Folta,et al.  Unexpected roles for cryptochrome 2 and phototropin revealed by high-resolution analysis of blue light-mediated hypocotyl growth inhibition. , 2001, The Plant journal : for cell and molecular biology.

[45]  C. Ballaré,et al.  Canopy Light and Plant Health1 , 2012, Plant Physiology.

[46]  C. Fankhauser,et al.  The Arabidopsis PHYTOCHROME KINASE SUBSTRATE2 Protein Is a Phototropin Signaling Element That Regulates Leaf Flattening and Leaf Positioning1[W][OA] , 2010, Plant Physiology.

[47]  S. Hoad,et al.  Effects of pre-severance light quality on the vegetative propagation of Eucalyptus grandis W. Hill ex Maiden. Cutting morphology, gas exchange and carbohydrate status during rooting. , 1996 .

[48]  Samuel Rossel,et al.  Navigation by bees using polarized skylight , 1993 .

[49]  R. Bula,et al.  Evaluation of light emitting diode characteristics for a space-based plant irradiation source. , 1992, Advances in space research : the official journal of the Committee on Space Research.

[50]  A. Simmons,et al.  LIME GREEN LIGHT‐EMITTING DIODE EQUIPPED YELLOW STICKY CARD TRAPS FOR MONITORING WHITEFLIES, APHIDS AND FUNGUS GNATS IN GREENHOUSES , 2004 .

[51]  Hong Wang,et al.  Effects of light quality on CO2 assimilation, chlorophyll-fluorescence quenching, expression of Calvin cycle genes and carbohydrate accumulation in Cucumis sativus. , 2009, Journal of photochemistry and photobiology. B, Biology.

[52]  A. Schuerger,et al.  Spectral quality affects disease development of three pathogens on hydroponically grown plants. , 1997, HortScience : a publication of the American Society for Horticultural Science.

[53]  M. Dorais,et al.  DEVELOPMENTAL AND PHYSIOLOGICAL RESPONSES OF TOMATO AND CUCUMBER TO ADDITIONAL BLUE LIGHT , 2006 .

[54]  G. Kennedy,et al.  Effect of day length and light intensity on 2-tridecanone levels and resistance inLycopersicon hirsutum f.glabratum toManduca sexta , 1981, Journal of Chemical Ecology.

[55]  K. Shoji,et al.  Effect of light quality on rosmarinic acid content and antioxidant activity of sweet basil, Ocimum basilicum L. , 2009 .

[56]  Gary W. Stutte,et al.  Photoregulation of Bioprotectant Content of Red Leaf Lettuce with Light-emitting Diodes , 2009 .

[57]  Hans Lambers,et al.  Plant Physiological Ecology , 2000, Springer New York.

[58]  B. A. Eveleens-Clark,et al.  Growth of tomatoes under hybrid led and HPS lighting , 2012 .

[59]  D. Fanourakis,et al.  Red and Blue Light Effects during Growth on Hydraulic and Stomatal Conductance in Leaves of Young Cucumber Plants , 2012 .

[60]  E. Huq,et al.  PHYTOCHROME-INTERACTING FACTOR 1 Is a Critical bHLH Regulator of Chlorophyll Biosynthesis , 2004, Science.

[61]  Cary A. Mitchell,et al.  Plant Productivity in Response to LED Lighting , 2008 .

[62]  A. Brazaitytė,et al.  LED illumination affects bioactive compounds in romaine baby leaf lettuce. , 2013, Journal of the science of food and agriculture.

[63]  M. Kirschbaum,et al.  Gas Exchange Analysis of the Relative Importance of Stomatal and Biochemical Factors in Photosynthetic Induction in Alocasia macrorrhiza. , 1988, Plant physiology.

[64]  Ahmad,et al.  Cryptochrome 1 controls tomato development in response to blue light. , 1999, The Plant journal : for cell and molecular biology.

[65]  T. Shinomura,et al.  Action spectra for phytochrome A- and B-specific photoinduction of seed germination in Arabidopsis thaliana. , 1996, Proceedings of the National Academy of Sciences of the United States of America.

[66]  J. Sullivan,et al.  Ultraviolet-B effects on stomatal density, water-use efficiency, and stable carbon isotope discrimination in four glasshouse-grown soybean (Glyicine max) cultivars , 2005 .

[67]  Diversity in plant red pigments: anthocyanins and betacyanins , 2013, Plant Biotechnology Reports.

[68]  R M Wheeler,et al.  Improving spinach, radish, and lettuce growth under red light-emitting diodes (LEDs) with blue light supplementation. , 2001, HortScience : a publication of the American Society for Horticultural Science.

[69]  Lydie Huché-Thélier,et al.  Light signaling and plant responses to blue and UV radiations—Perspectives for applications in horticulture , 2016 .

[70]  T. Dougher,et al.  Long-term Blue Light Effects on the Histology of Lettuce and Soybean Leaves and Stems , 2004 .

[71]  I. Kataoka,et al.  Effect of red- and blue-light-emitting diodes on growth and morphogenesis of grapes , 2008, Plant Cell, Tissue and Organ Culture.

[72]  H. Drumm-Herrel,et al.  Coaction between phytochrome and blue/UV light in anthocyanin synthesis in seedlings , 1983 .

[73]  J. J. Meyer,et al.  The phenolic, 3,4-dihydroxybenzoic acid, is an endogenous regulator of rooting in Protea cynaroides , 2007, Plant Growth Regulation.

[74]  Kazuhiro Fujiwara,et al.  Photosynthetic characteristics of rice leaves grown under red light with or without supplemental blue light. , 2004, Plant & cell physiology.

[75]  C. Brown,et al.  Life cycle experiments with Arabidopsis grown under red light-emitting diodes (LEDs). , 1998, Life support & biosphere science : international journal of earth space.

[76]  Gareth I. Jenkins,et al.  UV‐B Action Spectrum for UVR8‐Mediated HY5 Transcript Accumulation in Arabidopsis , 2009, Photochemistry and photobiology.

[77]  H. Watanabe,et al.  Effects of light quality on the growth and essential oil content in sweet basil , 2011 .

[78]  J. Casal,et al.  Cryptochrome as a Sensor of the Blue/Green Ratio of Natural Radiation in Arabidopsis1[C][W][OA] , 2010, Plant Physiology.

[79]  Cole Gilbert,et al.  Spectral efficiency of the western flower thrips, Frankliniella occidentalis , 1992 .

[80]  E. Wellmann,et al.  ANALYSIS OF LIGHT‐CONTROLLED ANTHOCYANIN FORMATION IN COLEOPTILES OF Zea mays L.: THE ROLE OF UV‐B, BLUE, RED AND FAR‐RED LIGHT , 1985 .

[81]  G. Holmes,et al.  Survival of Pseudoperonospora cubensis sporangia exposed to solar radiation , 2010 .

[82]  Jessica L. Gilbert,et al.  Light modulation of volatile organic compounds from petunia flowers and select fruits , 2013 .

[83]  T. Kozai,et al.  Sustainable plant factory: closed plant production systems with artificial light for high resource use efficiencies and quality produce , 2013 .

[84]  H. Watanabe,et al.  GROWTH OF BANANA PLANTLETS CULTURED IN VITRO UNDER RED AND BLUE LIGHT-EMITTING DIODE (LED) IRRADIATION SOURCE , 2002 .

[85]  D. Bell-Pedersen,et al.  Circadian Rhythms in Neurospora crassa and Other Filamentous Fungi , 2006, Eukaryotic Cell.

[86]  C. Kubota,et al.  Applications of far-red light emitting diodes in plant production under controlled environments , 2012 .

[87]  Andrei P. Sommer,et al.  Plants grow better if seeds see green , 2006, Naturwissenschaften.

[88]  K. Roh,et al.  Genes up-regulated during red coloration in UV-B irradiated lettuce leaves , 2007, Plant Cell Reports.

[89]  E. Tobin,et al.  Photobiological properties of the inhibition of etiolated Arabidopsis seedling growth by ultraviolet-B irradiation. , 2009, Plant, cell & environment.

[90]  A. Teramura,et al.  The Role of Flavonol Glycosides and Carotenoids in Protecting Soybean from Ultraviolet-B Damage , 1993, Plant physiology.

[91]  L. Corrochano Fungal photoreceptors: sensory molecules for fungal development and behaviour , 2007, Photochemical & photobiological sciences : Official journal of the European Photochemistry Association and the European Society for Photobiology.

[92]  Hyeon-Hye Kim,et al.  Green-light supplementation for enhanced lettuce growth under red- and blue-light-emitting diodes. , 2004, HortScience : a publication of the American Society for Horticultural Science.

[93]  Qichang Yang,et al.  Using Movable Light-emitting Diodes for Electricity Savings in a Plant Factory Growing Lettuce , 2014 .

[94]  M. Tanaka,et al.  Responses of strawberry plantlets cultured in vitro under superbright red and blue light-emitting diodes (LEDs) , 2003, Plant Cell, Tissue and Organ Culture.

[95]  Wei Fang,et al.  Effects of Frequency and Duty Ratio on the Growth of Potato Plantlets In Vitro Using Light-emitting Diodes , 2004 .

[96]  R. Tewari,et al.  Photon flux density and light quality induce changes in growth, stomatal development, photosynthesis and transpiration of Withania Somnifera (L.) Dunal. plantlets , 2007, Plant Cell, Tissue and Organ Culture.

[97]  Eiji Goto,et al.  EFFECTS OF BLUE AND RED LIGHT ON STEM ELONGATION AND FLOWERING OF TOMATO SEEDLINGS , 2012 .

[98]  A. Suthaparan,et al.  Specific Light-Emitting Diodes Can Suppress Sporulation of Podosphaera pannosa on Greenhouse Roses. , 2010, Plant disease.

[99]  J. Guiamet,et al.  Application of low intensity light pulses to delay postharvest senescence of Ocimum basilicum leaves , 2013 .

[100]  J. Chory,et al.  Phytochrome signaling mechanisms and the control of plant development. , 2011, Trends in cell biology.

[101]  G. Buck-Sorlin,et al.  Optimizing illumination in the greenhouse using a 3D model of tomato and a ray tracer , 2014, Front. Plant Sci..

[102]  A. Vian,et al.  Plant responses to red and far-red lights, applications in horticulture , 2016 .

[103]  A. Cashmore,et al.  Cryptochromes: enabling plants and animals to determine circadian time. , 2003, Cell.

[104]  A. Shimada,et al.  Red and blue pulse timing control for pulse width modulation light dimming of light emitting diodes for plant cultivation. , 2011, Journal of photochemistry and photobiology. B, Biology.

[105]  A. Manukyan Effects of PAR and UV‐B Radiation on Herbal Yield, Bioactive Compounds and Their Antioxidant Capacity of Some Medicinal Plants Under Controlled Environmental Conditions , 2013, Photochemistry and photobiology.

[106]  C. Brown,et al.  Photomorphogenesis, photosynthesis, and seed yield of wheat plants grown under red light-emitting diodes (LEDs) with and without supplemental blue lighting. , 1997, Journal of experimental botany.

[107]  Y. Honda,et al.  Light-induced Resistance of Broad Bean against Botrytis cinerea , 1998 .

[108]  Roberta Croce,et al.  Photosynthetic Quantum Yield Dynamics: From Photosystems to Leaves[W][OA] , 2012, Plant Cell.

[109]  N. Paul,et al.  Ultraviolet Radiation as a Limiting Factor in Leaf Expansion and Development , 2009, Photochemistry and photobiology.

[110]  X. Deng,et al.  Plant hormone signaling lightens up: integrators of light and hormones. , 2010, Current opinion in plant biology.

[111]  K. Mccree THE ACTION SPECTRUM, ABSORPTANCE AND QUANTUM YIELD OF PHOTOSYNTHESIS IN CROP PLANTS , 1971 .

[112]  Thomas C. Vogelmann,et al.  Green Light Drives CO2 Fixation Deep within Leaves , 1998 .

[113]  How-Chiun Wu,et al.  Red Light-emitting Diode Light Irradiation Improves Root and Leaf Formation in Difficult-to-propagate Protea cynaroides L. Plantlets In Vitro , 2012 .

[114]  Robert C. Morrow,et al.  Comparison of Intracanopy Light-emitting Diode Towers and Overhead High-pressure Sodium Lamps for Supplemental Lighting of Greenhouse-grown Tomatoes , 2013 .

[115]  Hendrik Poorter,et al.  Blue light dose–responses of leaf photosynthesis, morphology, and chemical composition of Cucumis sativus grown under different combinations of red and blue light , 2010, Journal of experimental botany.

[116]  H. Watanabe,et al.  In vitro growth of Cymbidium plantlets cultured under superbright red and blue light-emitting diodes (LEDs) , 1998 .

[117]  T. Sharkey,et al.  Light-emitting diodes as a light source for photosynthesis research , 2004, Photosynthesis Research.

[118]  R. Paradiso,et al.  Spectral dependence of photosynthesis and light absorptance in single leaves and canopy in rose , 2011 .

[119]  W. Backhaus Color vision and color choice behavior of the honey bee , 1993 .

[120]  Y. W. Kim,et al.  Growth of Tsuru-rindo (Tripterospermum japonicum) culturedin vitro under various sources of light-emitting diode (LED) irradiation , 2006, Journal of Plant Biology.

[121]  Zhi-gang Xu,et al.  The effects of different light qualities on rapeseed (Brassica napus L.) plantlet growth and morphogenesis in vitro , 2013 .