Horticultural Lighting Metrics

Looking Beyond Light Recipes

Ian Ashdown, P. Eng, FIES

Senior Scientist, Lighting Analysts Inc. / SunTracker Technologies Ltd.

[Please send all comments to allthingslighting@gmail.com]

UPDATE 17/08/26 – This article was first published on August 25th, 2017 in Urban Ag News.

It was all so easy until recently. Plants require light in order to grow, and so we provided them with daylight and/or electric lighting. Given the singular choice of high-pressure sodium (HPS) lamps, we only needed to be concerned about measuring Photosynthetically Active Radiation (PAR) and Daily Light Integrals (DLI).

The introduction of light-emitting diodes (LEDs) and solid-state lighting (SSL) has changed everything. With the ability to independently control the light source spectrum from ultraviolet through visible light to far-red, researchers and growers are discovering that plant species and even cultivars respond differently to different spectral power distributions. From these discoveries are coming “light recipes” for optimal plant growth and health.

Light recipes require more than a pinch of salt and a dash of cayenne, however. We need to measure and quantify the light received by plants, much as professional lighting designers have long measured and quantified light for building interiors and outdoor areas. These designers have numerous of metrics to call upon, all of them based on the human perception of visible light. Unfortunately, plants do not respond to light as we do, and so units of measurements such as lumens, lux, and candela are all but meaningless for horticultural lighting.

Given this, the American Society of Agricultural and Biological Engineers has just announced the publication of ANSI/ASABE S640 JUL 2017, Quantities and Units of Electromagnetic Radiation for Plants (Photosynthetic Organisms). Developed over two years by an international team of experts from industry and academia, this standard brings some much-needed order to the metrics of horticultural lighting.

The document formally defines 33 electromagnetic radiation metrics for horticultural lighting. They are fully compatible with metrics previously defined by standards from the American Society of Agricultural Engineers (precursor of the ASABE), the Illuminating Engineering Society (IES), the Commission Internationale d’Eclairage (CIE), and the International Organization for Standards (ISO). They are, however, specific to the needs of horticulture and plant biology.

What we perceive as visible light spans the electromagnetic spectrum with wavelengths from 400 nm (deep blue) to 700 nm (deep red). Coincidentally, this is the same range over which plant photosynthesis occurs. Outside of this range, plants respond to ultraviolet and far-red radiation. The Pfr isoform of phytochrome, for example, has a peak spectral absorptance of 735 nm, and is responsible for initiating many photomorphogenetic functions. Similarly, the photopigment UVR8 is responsible for sensing excess UV-B radiation (280 nm – 315 nm) and initiating plant stress responses to prevent DNA damage. With this, the metrics are therefore divided into three spectral ranges: ultraviolet (280 nm – 400 m), photosynthetic (400 nm – 700 nm), and far-red (700 nm – 800 nm).

The other division of the metrics is based on radiant versus photon flux. Every photon has a specific wavelength (e.g., 555 nm), and its energy (stated in watt-seconds, or joules) is inversely proportional to its wavelength. Plant photosynthesis does not care about photon energy, however – the chlorophyll molecule absorbs the photon for its chemical action and releases any excess energy as heat. Thus, horticulturalists and plant biologists are interested in the flow (or “photon flux”) of photons per second, with no regard for wavelength. This flux is measured in micromoles (6.23 ×1017) of photons per second with a broadband “quantum sensor,” typically a silicon photodiode with an optical filter.

Forest ecologists, on the other hand, are often interested in the energy of sunlight incident on the forest canopy, and so they measure electromagnetic radiation in terms of “radiant flux,” stated in watts. Here, wavelength matters, with blue light photons having more energy than red light photons. A broadband sensor, again typically a silicon photodiode with an optical filter, is used to measure radiant flux over the spectral range of interest.

It is also important to be able to measure and quantify the spectral power distribution of light sources with a spectroradiometer. In one recent study, for example, a difference of 10 nm in the peak wavelength of green LEDs (520 nm versus 530 nm) had a pronounced effect on the growth and development of red leaf lettuce (Johkan et al. 2012). We therefore have both spectral radiant flux and spectral photon flux, measured in watts per nanometer and micromoles per second per nanometer respectively.

With these divisions, we have the following horticultural lighting metrics defined by ANSI/ASABE S640:

Spectral Range Radiant Photon
 (280 nm – 800 nm) Flux Flux
Intensity Intensity
Efficiency Efficacy
Photosynthetic

(400 nm – 700 nm)

Flux Flux
Flux Density
Intensity Intensity
Efficiency Efficacy
Daily Light Integral
Ultraviolet

(280 nm – 400 nm)

Flux Flux
Flux Density
Intensity Intensity
Efficiency Efficacy
Far-red

(700 nm – 800 nm)

 

Flux Flux
Flux Density
Intensity Intensity
Efficiency Efficacy
Spectral (per nm) Flux Flux
Irradiance Flux Density
Intensity Intensity
Power Distribution Quantum Distribution

For now, horticulturalists will continue to measure PAR as photosynthetic photon flux density (PPFD) with a quantum sensor, and measure or calculate daily light integrals (integrated daily PPFD). However, ANSI/ASABE S640 is important in that it provides a framework with which to quantify forthcoming light recipes for optimal growth and health of urban agriculture crops.

Looking beyond light recipes, horticultural luminaire manufacturers will be able to quantify the optical performance characteristics of their products, and lighting design software developers will be able to develop products specifically for horticultural lighting design in greenhouses and vertical farms. It all begins, however, with horticultural lighting metrics.

ANSI/ASABE S640 is available for purchase from the ASABE Technical Library (https://elibrary.asabe.org).

References

Jokhan, M, et al. 2012. “Effect of Green Light Wavelength and Intensity on Photomorphogenesis and Photosynthesis in Lactuca sativa,” Environmental and Experimental Botany 75:128-133.

 

Phytochrome and PSS

Think Beyond Pink

Ian Ashdown, P. Eng., FIES

Senior Scientist, Lighting Analysts Inc. / SunTracker Technologies Ltd.

[Please send all comments to allthingslighting@gmail.com]

Horticultural lighting is currently one of the fastest-expanding markets in commercial lighting, with projected revenues of several billion dollars in less than a decade. From the perspective of a professional lighting designer, the market opportunities are enticing. Whether it is lighting for greenhouses or vertical farms and plant factories, the basic principles of lighting design remain the same.

FIG. 1 – Horticultural lighting in greenhouses. (Source: Colorado State University)

FIG. 1 – Horticultural lighting in greenhouses. (Source: Colorado State University)

There are, however, design metrics that will be unfamiliar to most lighting designers. One of these – the subject of this article – has the rather unwieldy name of phytochrome photostationary state (PSS). While rarely discussed outside of horticultural research publications, it represents an important concept for horticulturists, and particularly floriculturists.

To understand this metric, it is necessary to review some aspects of plant biology.

Photomorphogenesis

The development of plants, from seed to flowering, is very much dependent on the electromagnetic radiation they are exposed to. This developmental process, called photomorphogenesis, is completely separate from the process of photosynthesis. It relies on various photopigments, including phytochromes, cryptochromes, phototropins, and UVR8, to sense and respond to radiation ranging from ultraviolet to near-infrared.

Our interest is in the photopigment family of phytochromes, which are mostly sensitive to red and far-red visible radiation. They mediate the germination of seeds (photoblasty), the growth of stems and leaves toward visible light (etiolation), the time of flowering based on the length of day and night (photoperiodism), the synthesis of chlorophyll for photosynthesis, and more (e.g., Smith 2000). While there are six known members of the phytochrome family, it is convenient to refer to them generically and collectively as “phytochrome.”

Phytochrome exists in two states, or isoforms. In its ground state (identified as Pr), phytochrome strongly absorbs red light, and so appears turquoise-blue in concentrated solution in vitro (Figure 2). When it absorbs a red photon, however, it changes its physical shape to form its physiologically active state Pfr. In doing so, its peak spectral absorptance shifts towards the far-red, with a concentrated solution of phytochrome appearing more greenish in color.

When phytochrome is in its Pfr state, it may absorb a far-red photon and change once again into its Pr  state. This bistable behavior makes phytochrome an ideal biochemical switch, with the Pfr isoform serving as the “signaling” state to the plant. As one example of this biological function, red light typically penetrates several centimeters into loose soil (e.g., Borthwick et al. 1952, Botto et al. 1996). As the sun rises higher each day in the spring, an increasing amount of red light reaches the seeds, until a sufficient concentration of phytochrome switches from its Pr isoform to its Pfr isoform. This signals the cellular mechanisms of the seed that it is time to sprout. If, on the other hand, the seed is buried too deeply, it will never sprout and will eventually die.

FIG. 2 – Phytochrome spectral absorptance. (Source: Sager et al. 1988)

FIG. 2 – Phytochrome spectral absorptance. (Source: Sager et al. 1988)

Photoperiodism

The existence of phytochrome was first suspected nearly a century ago, when Garner and Allard (1920) studied the effects of day length on flowering plants. They observed that tobacco plants could be made to flower in summer by reducing the hours of daylight with artificial darkening, and that they could also be made to remain in a vegetative state during the winter by providing supplemental electric light. They called this effect photoperiodism.

Some plant species flower only when exposed to short periods of light (such as poinsettias – Islam et al. 2014), and so are called short-day plants, while others flower only after to exposure to long periods of light (such as spinach and radishes), and are called long-day plants. In some day-neutral plants (such as tomatoes), flowering is not regulated by photoperiod.

The reason for these reactions in both short-day and long-day plants is the response of phytochrome to red and far-red light. With short-day plants, exposure to a brief period of light during the night inhibits flowering, while the same exposure with long-day plants promotes flowering. Floriculturists can therefore use supplemental electric lighting to delay or advance the flowering of plants to meet market needs.

The traditional techniques for photoperiodic control include (Boyle 1992):

  • Increasing the day length

While high-pressure sodium (HPS) lamps have traditionally been used for supplemental greenhouse lighting, incandescent lamps have the advantage of being rich in red and far-red radiation. Compact and linear fluorescent lamps have also been used, but their relative lack of red and far-red radiation makes them ineffective for phytochrome response manipulation.

  • Night interruption

The phytochrome response to red and far-red radiation does not require continual exposure. Consequently, the flowering period can be influenced with only a few hours of electric lighting during the night. This has the advantage of being more energy-efficient.

  • Cyclic (intermittent) lighting

If incandescent rather than HPS lamps are used, it may be sufficient to pulse the lighting with a short duty factor, such as one minute every half hour. (The optimal duty factor will depend on the irradiance at the plant canopy.)

  • Shortening the day length

The plants are covered with an opaque material to reduce the irradiance, preferably to the equivalent of less than 20 lux of visible light. Typical materials are black sateen cloth, woven polyolefin sheeting, and black polyethylene films.

The problem with these techniques is that they are mostly trial-and-error with different plant species and greenhouse operation conditions. The goal is to manipulate the plant growth and development through the phytochrome response, but there is no practical means of quantitatively predicting the impact of these techniques.

Solid State Lighting

The introduction of solid state lighting to the horticultural industry has been nothing short of revolutionary. In addition to the energy savings afforded by the use of blue and red LEDs whose spectral power distributions (SPDs) are optimal for photosynthesis (Figure 1), the recent commercial availability of high-flux red and far-red LEDs from manufacturers such as Lumileds, Osram, and Cree means that horticulturists and floriculturists now have the ability to precisely tune the light source SPDs for optimal photoperiod control on a per-species basis.

While the LED manufacturers’ product names vary, the products of interest have peak spectral outputs at 660 nm and 730 nm, corresponding to the peak spectral absorptances of phytochrome isoforms Pr and Pfr, respectively (Figure 2). The key here is that the ratio of red to far-red light can be easily set or varied on a daily basis as required for photoperiodic control. Along with blue LEDs, this ability to precisely control the light source SPD leads to the promise of plant “light recipes,” where the SPD and other environmental factors can be chosen on a per-species basis, and possibly varied over the life cycle of the plant growth and development.

The problem, of course, is that in order to control something, you need to measure it. For professional lighting designers, you also need the ability to specify it.

PSS Metric

The phytochrome photostationary state (PSS) metric was introduced some two decades ago (Sager et al. 1998). It has been mostly of academic interest with HPS and incandescent lighting, but the introduction of LEDs for horticultural lighting has suddenly brought this metric to the forefront as a useful design tool.

The metric is conceptually simple: it is the ratio of the concentration of the Pr isoform of phytochrome to the total concentration of both isoforms:

phytochrome-and-pss-eqn-1    (1)

under constant irradiation by a light source. (The maximum value is less than unity because of the spectral overlap between the two isoforms.)

By itself, this seems of little to no interest to lighting designers – how do you measure the relative concentrations of the phytochrome isoforms in a plant? (It took nearly forty years from the time of Garner and Allard (1920) just to isolate phytochrome in the laboratory – Butler et al. 1959.)

What Sager and his fellow researchers did was to note that each phytochrome molecule could be conceptually modeled as an opaque sphere that fully absorbs any incident radiation. If you measure the spectral absorptance of the molecule in solution and know the concentration, you can calculate the equivalent photochemical cross-section of the molecule for each wavelength. (As an aside, LED manufacturers use exactly the same approach when modeling the optical characteristics of LED phosphors embedded in an epoxy or silicone matrix.)

With this, Sager et al. measured the spectral absorptance of the Pr and Pfr isoforms (reproduced in Figure 2) and expressed the results as phytochrome photochemical cross-sections, measured in square meters per mole (i.e., 6.022 x 1023 molecules), represented as sr and sfr, respectively. Equation (1) then becomes:

phytochrome-and-pss-eqn-2    (2)

where N(l) is the measured spectral photon flux for wavelength l over the range of 300 nm to 800nm.

For photometric test laboratories characterizing horticultural luminaires, all that needs to be done is to measure the luminaire’s relative spectral power distribution. Calculating the PSS metric using the photochemical cross-section data from Sager et al. (1988) in accordance with Equation 2 is then a simple spreadsheet exercise.

For professional lighting designers, it is even simpler: the PSS metric is a direct indication of the ability of the horticultural luminaire to manipulate the phytochrome isoforms. While this will also depend, of course, on the absolute irradiance at the plant canopy, the PSS metric reduces the spectral power distributions of the red and far-red LEDs to a single number that can be specified.

LED Color Binning

We are not done yet! Professional lighting designers are all too familiar with the issue of precision in lighting design metrics. For example, lamp and luminaire manufacturers typically report the CIE General Colour Rendering Index (CRI) using two digits, such as CRI = 92. If you refer to the history of the CRI metric’s development, however, you will learn that the intended precision of this metric is five units (van Trigt, 1999). In other words, the difference between CRI = 90 and CRI = 92 is visually imperceptible and so meaningless in terms of practical application.

The same question must be asked of the PSS metric. Is, for example, PSS = 0.39 quantitatively different from PSS = 0.38? Perhaps surprisingly, this is not a question for horticultural researchers. Rather, it is a question for lighting researchers and the lighting industry.

The underlying problem is a familiar one: LED color binning. Taking Lumiled’s LUXEON SunPlus 20 series of horticultural LEDs as typical examples, we have:

Table 1 – LUXEON SunPlus 20 peak wavelength binning. (Source: Lumileds 2016)

Product Name Peak Wavelength (nm)
Minimum Maximum
Deep Red 655 670
Far Red 720 750

The Lumileds product datasheet provides typical spectral power distributions for these two products with typical peak wavelengths, which can be digitized and shifted to represent the minimum, typical, and maximum peak wavelength SPDs, as shown in Figures 3 and 4. These figures also display the red Pr and far-red Pfr phytochrome spectral photochemical cross-sections (i.e., their spectral absorptances), with the SPDs normalized to the peak cross-sections for display purposes only.

FIG. 3 – Lumileds SunPlus 20 Deep Red

FIG. 3 – Lumileds SunPlus 20 Deep Red

FIG. 4 – Lumileds SunPlus 20 Far Red

FIG. 4 – Lumileds SunPlus 20 Far Red

From this information, and assuming that the peak spectral photon intensities of the red and far-red LEDs are the same, the phytochrome photostationary state (PSS) metric values can be calculated as follows:

Table 2 – Example PSS values range (2-nm resolution)

  Minimum Typical Maximum
PSS 0.3563 0.3797 0.4036

In other words, the PSS value may vary by approximately ±6 percent for a given luminaire manufacturer’s product. This is useful information for lighting designers when specifying or qualifying horticultural luminaire products, similar to the meaningful precision of the CRI metric.

There is one further question to address. Sager et al. (1988) reported the photochemical cross-section values sr and sfr at 2-nanometer intervals. This is useful from an academic perspective, but perhaps not so much from that of a photometric testing laboratory. Unless the laboratory performs the LED spectral power distribution measurements in-house, it is likely that the SPDs will be available in 5 nm increments only. While this data can be interpolated at 2-nm intervals for the purposes of calculating the PSS metric in accordance with Equation 2, will the difference in calculated results be significant?

To answer this question, the sr and sfr values published in Sager et al. (1988) were interpolated at 5-nm intervals using a cubic spline approximation, and Table 2 was recalculated using 5-nm resolution for the LED spectral power distributions:

Table 3 – Example PSS values range (5-nm resolution)

  Minimum Typical Maximum
PSS 0.3639 0.3873 0.4033

In this situation, the PSS value may vary by approximately ±5 percent for a given luminaire manufacturer’s product. More significantly, the PSS value for 5-nm resolution was only two percent higher than the PSS value with 2-nm resolution.

These results will of course vary for different typical PSS values, but likely not significantly.

As rules of thumb, therefore:

  • Differences in PSS values of less than ±5 percent are likely not significant.
  • It likely does not matter whether the PSS values are calculated using 2-nm or 5-nm resolution.

The qualifier “likely” recognizes that, while the PSS metric is some two decades old, greenhouse operators have yet to make use of it as a production tool. Future experience may indicate that these rules of thumb are too lax. In the meantime, however, the above analysis provides some guidance for both lighting designers and horticulturists.

Conclusion

Horticultural lighting presents interesting opportunities for professional lighting designers. It is a rapidly developing field where the use of blue and red LEDs for optimal photosynthesis is only the beginning. Solid state lighting has energized horticultural research into plant responses to light sources with different spectral power distributions, and there will surely be discoveries that improve our understanding of both photosynthesis and photomorphogenesis, as well as improvements in horticultural lighting design.

In the meantime, the phytochrome photostationary state (PSS) metric is an example of existing knowledge that will likely prove useful in designing and specifying horticultural lighting systems.

References

Borthwick, H.A., S. B. Hendricks, M. W. Parker, E. H. Toole, and V. K. Toole. 1952. “A Reversible Photoreaction Controlling Seed Germination,” Proceedings of the National Academy of Science 38:662–666.

Botto, J. F., R. A. Sánchez, G. C. Whitelam, and J. J. Casal. 1996. “Phytochrome A Mediates the Promotion of Seed Germination by Very Low Fluences of Light and Canopy Shade Light in Arabidopsis,” Plant Physiology 110:439-444.

Butler, W. L., K. H. Norris, H. W. Siegelman, and S. B. Hendricks. 1959. “Detection, Assay, and Preliminary Purification of the Pigment Controlling Photoresponsive Development of Plants,” Proc. National Academy of Sciences 45:1703-1708.

Boyle, T. 1992. “Photoperiod Control Systems for Greenhouse Crops,” Floral Notes 4(6):2-4. (See also Photoperiod Control Systems for Greenhouse Crops.)

Garner, W. W., and H. A. Allard. 1920. “Effect of the Relative Length of Day and Night and Other Factors of the Environment on Growth and Reproduction in Plants,” Journal of Agricultural Research 18:553-606.

Islam, M. A., D. Tarkowská, J. L. Clarke, D.-R. Blystad, H. R. Gislerød, S. Torre, and J. E. Olsen. 2014. “Impact of End-of-day Red and Far-red Light on Plant Morphology and Hormone Physiology of Poinsettia,” Scientia Horticulturae 174:77-86.

Lumileds. 2016. DS171 LUXEON SunPlus Series for Horticulture, Lumileds Product Datasheet 20160908.

Sager, J. C., W. O. Smith, J. L. Edwards, and K. L. Cyr. 1988. “Photosynthetic Efficiency and Phytochrome Equilibria Determination Using Spectral Data,” Trans. ASAE 31(5):1882-1889.

Smith, H. 2000. “Phytochromes and Light Signal Perception by Plants – An Emerging Synthesis,” Nature 407:585-591.

van Trigt, C. 1999. “Color Rendering, A Reassessment.” Color Research & Application 24(3):197-206.

 

Botanical Light Pollution

Red is the New Blue

Ian Ashdown, P. Eng., FIES

Chief Scientist, Lighting Analysts Inc.

[ Please send all comments to allthingslighting@gmail.com ]

Related Posts

Color Temperature and Outdoor Lighting

Light Pollution and Uplight Ratings

Mobile Light Pollution

Blue-rich light from LED streetlights, we are told, is the nemesis of professional and amateur astronomers. Blue light is preferentially scattered by the atmosphere, resulting in potentially unacceptable levels of light pollution for astronomical observations. Unfortunately, LED streetlights emit more blue light on a per-lumen basis than the high-pressure sodium streetlights they are rapidly replacing.

Botanists and horticulturalists, however, may choose to differ. For them, it is red light from streetlights that is the problem. Depending on the species and various environmental factors, even low levels of light trespass from roadway and outdoor area luminaires can have harmful effects on both wild and domesticated plants. LED streetlights likewise emit more red light on a per-lumen basis than high-pressure sodium streetlights.

This is not a newly discovered problem. Botanists were aware of the deleterious effects of incandescent street lighting on trees eighty years ago (Matske 1936), while horticulturalists became aware of the problem with respect to ornamental plants some forty years ago (Cathey and Campbell 1975).

The lighting community can perhaps be excused for not following the latest research in publications such as American Journal of Botany and Journal of Arboriculture, but we were in fact made aware of the issue through publication of an article in Lighting Design and Application (Cathey and Campbell 1974). However, given that the proposed solution then was to avoid using high-pressure sodium (HPS) lamps and instead use less-efficient mercury-vapor lamps with their ghoulish color rendering capabilities … well, we understandably ignored the advice.

Soybeans and Trees

This is not to say that farmers are not aware of the problem. If you are growing soybeans, you quickly learn not to plant them in a field adjacent to HPS roadway lighting (FIG.1). The nighttime illumination – even as little as two to eight lux – can reduce crop yield by 20 to 40 percent due to delayed flowering and ripening (Chen et al. 2009).

Botanical Light Pollution - FIG. 1FIG. 1 – Effect of light pollution on soybean crop. (Source: Chen et al. 2009)

Landscape designers and arborists are also aware of the problem. A publication from Purdue University, for example, lists 65 trees and shrubs that are vulnerable to artificial light (Chaney 2002). Exposure to nighttime illumination, particularly from HPS street lighting, may result in disruption of the plant’s shoot growth, flowering, leaf expansion and abscission, and bud dormancy. In temperate climates, this may make the plants more susceptible to frost, fungal infections, and insect infestations. Again, however, the advice was to avoid using HPS lighting and use mercury vapor lighting instead. For lighting designers, this is pointless advice – mercury vapor lamps were long ago replaced by high-pressure sodium lamps, and these in turn are being replaced by solid state lighting.

… and herein lies today’s issue. LED-based outdoor lighting may – and the emphasis is on the word may – exacerbate the problem from the perspective of wild and domesticated plants. High-pressure sodium lamps emit much more red light than mercury vapor lamps on a per-lumen basis, and white light LEDs may (depending on their correlated color temperature) emit even more. What was once a minor problem for landscape designers and urban arborists may become something that lighting designers will need to consider.

To better understand this issue, we first need to understand the role of photopigments in plant growth and development.

Phytochrome

Plants perform their magic of photosynthesis using a photopigment called chlorophyll, but this is only one of many different photopigments plants use to harvest and detect light. Equally important is phytochrome, which regulates a long list of plant functions, including:

  • Seed germination and development
  • Stem elongation
  • Leaf expansion and abscission
  • Photosynthesis development
  • Flowering
  • Ripening
  • Dormancy

Taken together, these functions basically outline the life cycle from seed to adult plant.

The sum of these light-induced changes is called photomorphogenesis. There are other photopigments involved, including blue light-sensitive cryptochromes (Lin 2002) and ultraviolet-sensitive UVR8 (Goto et al. 2006, Kami et al. 2010). However, it is phytochrome that dominates plant growth and development.

Phytochrome itself is an interesting pigment in that it has two states (or isoforms) called Pr and Pfr (e.g., Smith 2000). The Pr isoform strongly absorbs red light, with a spectral peak at about 660 nm (FIG. 2), making it look turquoise-blue when dissolved in solution. This is its biologically inactive state.

When a phytochrome molecule absorbs a red photon, it switches to its Pfr isoform, making it look slightly more greenish. This is its biologically active state, which signals to the plant that red light has been sensed. While in this state, phytochrome has a different spectral absorption distribution (FIG. 2), with a spectral peak at about 730 nm. (Horticulturalists and plant biologists refer to the spectral range of 700 nm to 800 nm as “far-red,” which explains the “fr” subscript.)

When the Pfr isoform absorbs a far-red photon, it reverts to its Pr isoform. Thus, phytochrome performs the function of a resettable biological switch to initiate or terminate photomorphological processes.

Botanical Light Pollution - FIG. 2

FIG. 2 – Phytochrome absorption spectra. (Source: Plants in Action, First Edition[1])

This biological switch behavior has some interesting consequences. While even low levels of red light can initiate many physiological responses, applying far-red light soon thereafter may reset the switch and terminate the response. Light pulses as short as one minute at night – think car headlights on a country road – are enough to induce or prevent the flowering of some plants (Borthwick et al. 1952). Worse, some plants have flower induction thresholds of less than four lux (Botto et al. 1996, Whitman et al. 1998).

Photoperiodism

Phytochrome may act as a biological switch, but how plants respond to its signaling varies by species and even cultivar. What all plants have in common, however, is photoperiodism, their physiological reaction to the length of the day. Like humans and all other animals, plants have circadian rhythms.

In terms of flowering, plants can generally be divided into three categories: 1) short day; 2) long day; and 3) day-neutral. For short day plants, flowering is initiated, advanced, or promoted when the dark nighttime period is sufficiently long to allow enough phytochrome Pfr to revert to Pr. For long day plants, the opposite is true: flowering is initiated, advanced, or promoted when the dark nighttime period is sufficiently short to increase nighttime levels of phytochrome Pr. As for day-neutral plants, their time of flowering is determined by other environmental cues, such as temperature and moisture.

From the perspective of wild and domesticated plants growing outdoors, artificial light can be a problem. For horticulturalists, however, it can be a boon. Florists have long used incandescent lamps with their copious red and infrared emissions to modify the growth and development of flowering plants in greenhouses. This promotes flowering in long day plants such as asters, azaleas, and fuchsias, while delaying flowering in short day plants such as chrysanthemums, begonias, and poinsettias.

The recent availability of high-power red and far-red LEDs has provided new opportunities for both florists and horticulturalists. Independently switching or dimming these LEDs enables greenhouse operators to precisely control phytochrome as a biological switch. This, combined with the secondary effects of activating cryptochromes using blue light, provides remarkable control of plant growth and development (e.g., Gautam et al. 2015, Islam et al. 2014, Kitazaki et al. 2015, and Lee et al. 2015).

Light Pollution

Outside of the greenhouse environment, however, adding red and far-red radiation to the environment is not a good thing. We can call it what it is: botanical light pollution. For soybean farmers and urban arborists, it may be a nuisance. However, there can also be more insidious and detrimental effects for wild plants and the pollinating insects that depend on them (e.g., Bennie et al. 2016).

The question is, how do we quantify this pollution? It is reasonably easy to quantify astronomical light pollution because we have comprehensive mathematical models of atmospheric physics and optics. However, the best that botanists can do for us is to identify plants as short day, long day, or day neutral.

Pragmatically speaking, we do not need to quantify botanical light pollution in an absolute sense of so many micromoles of radiation per square meter per second or whatever. From a lighting design perspective, the goal is to illuminate an area with so many lumens per square meter while doing our best to prevent wasted spill light. The question then becomes, what is the best light source for plants?

Comparing Light Sources

The phytochrome absorptance spectra (FIG. 2) were obtained by extracting phytochrome from plants and dissolving it in solution for analysis in vitro with a spectrophotometer. When in the plant itself, however, phytochrome is surrounded by other photopigments, especially chlorophyll. Both chlorophyll A and chlorophyll B have absorptance spectra that overlap with those of the phytochrome isoforms (FIG. 3), so it is reasonable to ask whether this influences (or “screens”) the phytochrome absorptance spectra in vivo.

Botanical Light Pollution - FIG. 3

FIG. 3 – Photopigment spectral absorptances.

Fortunately, a variety of studies of the effect of monochromatic radiation on plant growth and development have shown that the absorptance spectra of phytochrome in vitro reasonably predict the plant physiological response. For example, Withrow et al. (1957) studied the “induction and reversion of hypocotyl hook opening” in bean seedlings. A plot of their results as induction and reversion “action spectra” shows a remarkable correlation with the in vitro absorptance spectra of phytochrome (FIG. 4).

Botanical Light Pollution - FIG. 4

FIG. 4 – Typical phytochrome action spectra. (Source: Smith 1977)

Given this, we can use the phytochrome absorptance spectra as a species-independent measure of the effect of red and far-red radiation on plant growth and development (Sager et al. 1988). For a given light source, the probability of a phytochrome molecule absorbing a photon with a given wavelength is determined by the absorptance spectra of the isoform and the relative number of photons with that wavelength.

For a light source, we typically have its relative spectral power distribution (SPD), which is measured in watts per nanometer. However, from the Planck-Einstein relation, we know that a photon’s energy is inversely proportional to its wavelength. Therefore, to determine the relative spectral photon flux distribution, we need only multiply the lamp SPD by the wavelength for each wavelength and normalize the resultant graph. (An example is shown in FIG. 5.)

Botanical Light Pollution - FIG. 5

FIG. 5 – Radiant versus photon flux for a 3000K warm white LED.

With this, we now have the means to compare light sources with different spectral power distributions. Given a reference lamp (say, HPS) and a test lamp (say, a 3000K warm white LED), the calculations consist of:

  1. Multiply the SPD values of each lamp by the CIE 1931 luminous efficiency function V(l) shown in FIG. 6 from 400 nm to 700 nm.
  2. Sum the results of Step 1 to obtain the relative lumens Fref and Ftest generated by the two lamps.
  3. Multiply the SPD values of the test lamp by Fref / Ftest.

The two SPDs now represent the same number of photopic lumens (i.e., luminous flux) emitted by the lamps. With this:

  1. Multiply the SPD values of each lamp by the wavelength to obtain the lamp spectral photon flux distributions from 500 nm to 800 nm.
  2. Multiply the results of Step 4 by the phytochrome Pr spectral absorptance spectrum.
  3. Sum the results of Step 5 to obtain the Pr action values pAref,r and pAtest,r.
  4. Multiply the results of Step 4 by the phytochrome Pfr spectral absorptance spectrum.
  5. Sum the results of Step 7 to obtain the Pfr action values pAref,fr and pAtest,fr.

and finally:

  1. Add the Pr and Pfr action values for each lamp to obtain the lamp phytochrome action values pAref and pAtest.
  2. Divide pAtest by pAref to obtain the relative action value for the test lamp compared to the reference lamp.

Botanical Light Pollution - FIG. 6

FIG. 6 – CIE 1931 luminous efficiency function V(l).

A few explanatory notes:

  1. As shown in FIG. 2, the phytochrome absorptance spectra have secondary peaks in the near-ultraviolet. These are ignored because: a) it is difficult to disentangle the effects of phytochrome from the effects of the blue-sensitive cryptochrome photopigments; and 2) the photomorphological effects of blue light are less pronounced than those resulting from red and far-red radiation. The lower limit of 500 nm was chosen based on the phytochrome absorptance spectra minima.
  2. The spectral peak of Pfr is only 60 percent that of Pr, but the area under each spectral curve between 500 nm and 800 nm is almost the same. Also, phytochrome action spectra for various plant species have shown that equal red and far-red radiant fluences at the spectral peaks of 660 nm and 730 nm have approximately equal effect on the physiological responses. This justifies the final step of adding the two action values.

It must be emphasized that these “action values” are approximate at best, and should not be considered as formally quantifiable metrics. They are introduced here only to explore the potential effects of botanical light pollution.

With this caveat then, the following light sources were selected for comparison:

Light Source Manufacturer Product Code
High-pressure sodium (test) Damar 1782 LU100M
2700K white light LED Lumileds LUXEON Rebel ES LXW9-PW27
3000K white light LED Lumileds LUXEON Rebel ES LXW9-PW30
3500K white light LED Lumileds LUXEON Rebel ES LXW8-PW35
4000K white light LED Lumileds LUXEON Rebel ES LXH7-PW40
5000K white light LED Lumileds LUXEON Rebel ES LXW8-PW40

Table 1 – Comparison light sources.

The HPS lamp SPD was measured in the laboratory with 0.1 nm resolution and averaged to 5 nm bins, while the Lumileds SPDs were digitized from the published datasheet (Lumileds 2014). The equal-lumen SPDs for these light sources are shown in FIG. 7.

Botanical Light Pollution - FIG. 7

FIG. 7 – Equal-lumen spectral power distributions.

Following the above calculation procedure with the HPS lamp as the test source, the relative phytochrome action values are:

Light Source Relative Phytochrome Action
Pr Pfr Pr + Pfr
High-pressure sodium 1.0 1.0 1.0
2700K white light LED 1.7 2.3 1.9
3000K white light LED 1.5 2.0 1.7
3500K white light LED 1.0 1.2 1.1
4000K white light LED 1.0 1.0 1.0
5000K white light LED 0.9 1.0 0.9

Table 2 – Relative phytochrome action values.

From this, it can be seen that while 2700K and 3000K white light LEDs produce the least astronomical light pollution (see related article Color Temperature and Outdoor Lighting), they also unfortunately produce the most botanical light pollution.

It should be noted however that these results apply to Lumileds LUXEON products only. Looking at FIG. 7, it is evident that the 2700K and 3000K products use a different phosphor formulation than the 3500K, 4000K, and 5000K products. Different major LED manufacturers will have their own proprietary phosphor formulations, and so the above results should not be applied to LEDs based solely on their nominal CCTs.

Add More Red

It seems counterintuitive, but one solution to the problem of excess red light generated by low-CCT LEDs is to add more red light.

Some of the early LED modules combined phosphor-coated white and red LED dice in order to compensate for the low-efficiency red phosphors then available. This produced a warm white light with good CIE Ra values, but relatively poor R9 values due to the quasimonochromatic red emissions.

One roadway luminaire manufacturer has recently taken this approach with a new product line that was reportedly designed to comply with the International Dark Sky Association’s Fixture Seal of Approval program requirements for a maximum CCT of 3000K. While the approach works (with a measured CCT of 3145K), the massive spike in red light peaking at 625 nm (see Fig. 8) would seem to be a botanist’s nightmare spectrum.

Botanical Light Pollution - FIG. 8

FIG. 9 – 3000K LED versus 3145K white+red LED equal-lumen spectral power distribution.

Surprisingly, the situation may not be as bad as it appears. First, there is relatively little far-red radiation being emitted. Second, the 625 nm peak occurs where the phytochrome Pr absorptance spectrum is only 50 percent of maximum. This results in a calculated phytochrome action value (relative to the HPS reference lamp) of 0.9 – half that of the 3000K LED.

Light Source Relative Phytochrome Action
Pr Pfr Pr + Pfr
White+red LED 0.9 0.9 0.9

Table 3 – White+red relative phytochrome action values.

Color Filters

Another solution to the problem of excess red light is simply to add a color filter with a sharp cutoff at 625 nm. Red light beyond the cutoff wavelength contributes only ten percent to the luminous flux of a 3000K white light LED, so it is may be a reasonable tradeoff. (The resultant color will, however, be slightly cyan in hue.)

Whether it is possible to develop a suitable dye or coating for the LED optics that is both inexpensive and resistant to fading is, of course, an open question.

Chlorophyll Screening

The preceding analysis necessarily assumes that the phytochrome is not screened by the other plant photopigments, and that the isoform absorptance spectra represent the phytochrome action spectra for any given plant. In practice, this is not necessarily true. Phytochrome is present in very low concentrations in plant tissues. As a result, the much higher concentrations of chlorophyll tend to screen phytochrome by absorbing much of the incident red radiation. (See Fig. 3 for spectral overlapping between phytochrome Pr and chlorophyll A.)

A study by Beggs et al. (1980) demonstrated that if mustard seedlings are treated with the herbicide Norflurazon, the chlorophyll in the plant tissue becomes photobleached, resulting in white rather than green seedlings. With white seedlings, the phytochrome action spectrum had a peak at 660 nm, following the phytochrome Pr absorptance spectrum. With untreated green seedlings, however, the action spectrum was shifted to approximately 630 nm – which is well within the range of the 625 nm LED emission of the white+red LEDs (FIG. 9).

Botanical Light Pollution - FIG. 9

FIG. 9 – Chlorophyll screening of phytochrome Pr action spectrum. (Source: Beggs et al. 1980)

Summary

First and foremost, the phytochrome action metric presented in this article is not intended as a formal light source metric in any sense; it was introduced solely as a means of evaluating the potential impact of red and far-red light on both wild and domestic plants.

Second, the effects of applying red and/or far-red radiation will depend on the physiological state of the plant, the physiological response being mediated, and the time of application. Any excess (i.e., artificial) red radiation will convert the Pr isoform in the exposed plant to Pfr , while any excess far-red radiation will convert the Pfr isoform to Pr. Either action will upset the plant’s phytochrome photostationary state (Sager et al. 1988). What effect this will have on a given plant species at any given time of the night and season is unknown.

While phytochrome may function as a biological switch for plants, how individual plants species respond to its signaling will vary. Given that phytochrome mediates so many plant functions, the botanist’s characterization of short day, long day, and day neutral flowering plants is probably about all they will have in common.

If the above analysis has shown anything, it is that by changing roadway and outdoor area lighting from high-pressure sodium to white light LEDs, we may – and again, the emphasis is on may – be upsetting the ecological balance in unexpected ways. By examining what we do know and applying it on a theoretical basis, we can at least be better prepared to respond in the future if we need to.

References

Beggs, C. J., M. G. Holmes, M. Jabben, and E. Schäfer. 1980. “Action Spectra for the Inhibition of Hypocotyl Growth by Continuous Irradiation in Light and Dark-grown Sinapis alba L. Seedlings,” Plant Physiology 66:615-618.

Bennie, J. T. W. Davies, D. Cruse, and K. J. Gaston. 2016. “Ecological Effects of Artificial Light on Wild Plants,” Journal of Ecology (in press).

Borthwick, H.A., S. B. Hendricks, M. W. Parker, E. H. Toole, and V. K. Toole. 1952. “A Reversible Photoreaction Controlling Seed Germination,” Proceedings of the National Academy of Science 38:662–666.

Botto, J. F., R. A. Sánchez, G. C. Whitelam, and J. J. Casal. 1996. “Phytochrome A Mediates the Promotion of Seed Germination by Very Low Fluences of Light and Canopy Shade Light in Arasbidopsis,” Plant Physiology 110:439-444.

Cathey, H. M., and L. E. Campbell. 1974. “Lamps and Lighting – A Horticultural View,” Lighting Design and Application 4(11):41-52.

Cathey, H. M., and L. E. Campbell. 1975. “Security Lighting and its Impact on the Landscape,” Journal of Arboriculture 1(10):181-187.

Chaney, W. R. 2002. Does Night Lighting Harm Trees? Circular FNR-FAQ-17, Department of Forestry and Natural Resources, Purdue University.

Chen, C.-L., Y.-H. Su, C.-J. Liu, and Y.-C. Lee. 2009. “Effect of Night Illumination on Growth and Yield of Soybean,” Journal of Taiwan Agricultural Research 58(2):146-154.

Gautam, P., M. T. Terfa, J. E. Olsen, and S. Torre. 2015. “Red and Blue Light Effects on Morphology and Flowering of Petunia x hybrid,” Scientia Horticulturae 184:171-178.

Goto, N., T. Kumagai, and M. Koornneef. 2006. “Flowering Responses to Light-breaks in Photomorphogenic Mutants of Arabidopsis thaliana, a Long-day Plant,” Physiologia Plantarum 83(2):209-215.

Islam, M. A., D. Tarkowská, J. L. Clarke, D.-R. Blystad, H. R. Gislerød, S. Torre, and J. E. Olsen. 2014. “Impact of End-of-day Red and Far-red Light on Plant Morphology and Hormone Physiology of Poinsetta,” Scientia Horticulturae 174:77-86.

Kami, C., S. Lorrain, P. Hornitschek, and C. Fankhauser. 2010. “Light-regulated Plant Growth and Development,” Current Topics in Developmental Biology 91:29-66.

Kitazaki, K., S. Watanabe, A. Okamoto, M. Matsuo, S. Furuya, and K. Sameshima. 2015. “Far-red Light Enhances Removal of Pericarps in Tartary Buckwheat (Fagopyrum tataricum Gaertn.) Sprout Production under Artificial Lighting,” Scientia Horticulturae 185:167-174.

Lee, M.-J., S.-Y. Park, and M.-M. Oh. 2015. “Growth and Cell Division of Lettuce Plants under Various Ratios of Red to Far-red Light-emitting Diodes,” Horticulture, Environment, and Biotechnology 56(2):186-194.

Lin, C. 2002. “Blue Light Receptors and Signal Transduction,” The Plant Cell 14:S205-S225.

Lumileds. 2014. LUXEON Rebel ES Datasheet DS61 20140630.

Matzke, E. B. 1936. “The Effect of Street Lights in Delaying Leaf-fall in Certain Trees,” American Journal of Botany 23(6):446-452.

Sager, J. C., W. O. Smith, J. L. Edwards, and K. L. Cyr. 1988. “Photosynthetic Efficiency and Phytochrome Equilibria Determination Using Spectral Data,” Trans. ASAE 31(5):1882-1889.

Smith, H., Ed. 1977. The Molecular Biology of Plant Cells. Berkely, CA: University of California Press.

Smith, H. 2000. “Phytochromes and Light Signal Perception by Plants – An Emerging Synthesis,” Nature 407:585-591.

Whitman, C.M., R. D. Heins, A. C. Cameron, and W. H. Carlson. 1998. “Lamp Type and Irradiance Level for Daylength Extensions Influence Flowering of Campanula carpatica ‘Blue Clips’, Coreopsis grandiflora ‘Early Sunrise’, and Coreopsis verticillata ‘Moonbeam’,” Journal of the American Society of Horticultural Science 123:802–807.

Withrow, R. B., W. H. Klein, and V. Elstad. 1957. “Action Spectra of Photomorphogenic Induction and its Photoinactivation,” Plant Physiology 32:453-462

 

[1] Plants in Action is a plant physiology textbook published by the Australian Society of Plant Scientists, New Zealand Society of Plant Biologists, and the New Zealand Institute of Agricultural and Horticultural Science. It is freely available online at http://plantsinaction.science.uq.edu.au.

Photometry and Photosynthesis

From Illuminance to PPFD

Ian Ashdown, FIES

Chief Scientist, Lighting Analysts, Inc.

[ Please send comments to allthingslighting@gmail.com ]

Related Posts

Botanical Light Pollution

UPDATE 15/04/13 – This article was first published on December 10, 2014. A revised version was published as “LED Lighting for Horticulture” in the Mar/Apr 2015 issue of LED Professional Review (www.led-professional.com). This update includes information from the published article.

UPDATE 15/11/05 – due to several Excel spreadsheet errors, the lux-to-PPFD conversion factors presented in Table 2 were miscalculated. These errors have been corrected.

UPDATE 16/01/12 – the description of the Emerson effect has been corrected.

UPDATE 16/02/10 – Added discussion of calculating lumen-to-PPFD conversion factors for overcast skies, as well as expanded notes and references on green and ultraviolet LEDs.

Horticultural lighting these days is big business. As agricultural land becomes scarce, the weather becomes more unreliable, and the migration of people to megacities continues, it increasingly makes economic sense to cultivate plants indoors with electric lighting.

The problem is that lighting designers and horticulturalists generally do not speak the same language. Whereas we speak in terms of lumens and illuminance, horticulturalists speak in terms of photosynthetically active radiation (PAR) and photosynthetic photon flux density (PPFD). Ask for an explanation of these terms and you will hear talk of micromoles, and possibly microeinsteins, of photons. Instead of luminous flux, there is quantum flux. It can be very confusing, not to say frustrating.

We need however to understand each other. As the horticultural industry transitions from high-intensity discharge arc lamps such as high-pressure sodium (HPS) and metal halide (MH) to solid-state lighting (e.g., Massa et al. 2008, Mitchell et al. 2012, Nelson and Bugbee 2014), it becomes possible to tailor the spectral power distribution of multicolor-LED luminaires for individual crops and plant species. For lighting designers working with horticulturalists, the need to understand PAR and calculate PPFD values using lighting design software becomes critical.

Photosynthetically Active Radiation

Photosynthesis is the process used by plants to convert electromagnetic radiation – light – into chemical energy that is used for growth and development. All that is needed for this process is carbon dioxide (CO2), nutrients, and water. The process itself is not particularly efficient; only 4 to 6 percent of the absorbed radiation is converted into chemical energy (Zhu et al. 2010, Table 2). Still, it is the engine that drives most life on this planet.

Photosynthetically active radiation (PAR) is defined as electromagnetic radiation over the spectral range of 400 nm to 700 nm that photosynthetic organisms are able to use in the process of photosynthesis to fix the carbon in CO­2 into carbohydrates. Horticulturalists measure PAR for both plant research and greenhouse lighting design (e.g., Barnes et al. 1993) using specialized photometers (e.g., Biggs et al. 1971).

A common unit of measurement for PAR is photosynthetic photon flux density (PPFD), measured in units of moles per square meter per second[1]. In this case, every absorbed photon, regardless of its wavelength (and hence energy), is assumed to contribute equally to the photosynthetic process. This is in accordance with the Stark-Einstein law, which states that every photon (or quantum) that is absorbed will excite one electron, regardless of the photon’s energy, between 400 nm and 700 nm. For this reason, photosynthetic photon flux is also referred to as quantum flux.

Whether a photon with a given wavelength is absorbed by a plant leaf is dependent on the spectral absorptance of the leaf, which in turn is determined largely by the leaf optical properties, including the concentration of plant pigments such as chlorophyll A and B, various cartrenoids (carotenes and xanthophylls), and anthocyanins. The chlorophylls are responsible for the characteristic green color of leaves; the other pigments contribute to the yellow, orange, and red colors respectively of autumn leaves after the chlorophylls decompose.

Typical absorptance spectra for chlorophyll A, chlorophyll B, beta-carotene, and two isoforms of phytochrome are shown in Figure 1. It must be noted, however, that these spectra are approximate. They are measured in vitro by dissolving the pigments as extracts in a solvent, which affects their absorptance spectra. By themselves, they suggest that blue and red LEDs alone are sufficient for horticultural applications. In reality, however, the situation is much more complicated.

Photosynthesis - FIG 1 (Update)

FIG. 1 – Photopigment spectral absorptances

McCree (1972a) measured the spectral absorptance (FIG. 2) and quantum yield of CO2 assimilation for the leaves of 22 species of crop plants[2]. Taking the average measurements at 25 nm intervals for all plant species (Table 1), he produced the photon-weighted relative quantum yield (Table 1) that is representative of most crop plants.

Photosynthesis - FIG 1

FIG. 2 – Average crop plant spectral absorptance

Wavelength (nm) Relative Spectral Quantum Yield
400 0.42
425 0.68
450 0.70
475 0.63
500 0.65
525 0.72
550 0.82
575 0.91
600 0.97
625 1.00
650 0.90
675 0.90
700 0.48

Table 1 – Relative Quantum Yield (average of 22 field species)

Yield Photon Flux

It is also possible to measure PAR in terms of energy rather than photons. The energy of a photon with wavelength λ is given by the Planck-Einstein relation:

E = hc / λ

where E is the energy in joules, h is Planck’s constant (6.626 x 10-34 joule-seconds), c is the speed of light (2.998 x 108 meters per second), and l is measured in meters. For example, one micromole of photons with a wavelength of 450 nm has 0.266 joules of energy. Scaling the photon-weighted relative quantum yield values by the wavelength and normalizing produces the energy-weighted relative quantum yield, also known as the action spectrum (FIG. 3). (An action spectrum is simply a plot of biological effectiveness as a function of wavelength of incident light.)

The energy-weighted photosynthetic photon flux is measured in watts (joules per second), and is referred to as the yield photon flux (YPF). (In terms of lighting design, it is synonymous with irradiance.) As shown by McCree (1972b), photon-weighted PPFD is a better predictor of photosynthesis when light sources with different spectral power distributions are considered. Given this, PPFD is the metric most commonly used by horticulturalists. However, energy-weighted YPFD is useful for energy-balance calculations involving photosynthetic organisms.

Photosynthesis - FIG 2

FIG. 3 – Relative quantum yield for crop plant photosynthesis

As noted by McCree (1972a), neither PPFD (quantum flux density) nor YPFD (irradiance) are perfect measures of photosynthetically active radiation in that both systematically overestimate the effectiveness of blue light relative to red. As can be seen from FIG. 3, the error is greater for YPFD, which explains why PPFD measurements are preferred by horticulturalists. Nevertheless, they are useful in that they are independent of any particular plant species, and they can be measured both in the laboratory and in the field using a radiometer with a spectrally-calibrated quantum sensor such as the LI-190SA with LI-COR (www.licor.com).

From Lumens to Photosynthetic Photon Flux

As lighting designers, we need some method of converting lumens to quantum flux and illuminance to quantum flux density (PPFD). We can do so however only if we know or can estimate the spectral power distribution (SPD) of the light source.

Suppose then that we have a light source with a known relative spectral power distribution (SPD), such as for example a 5000K “cool white” LED (FIG. 4).

Photosynthesis - FIG 3

FIG. 4 – 5000K LED relative spectral power distribution

One watt of radiant power at 555 nm is by definition equal to 683 lumens. Given the CIE 1931 luminous efficiency function (FIG. 5), we can calculate the spectral radiant flux Φ(λ) in watts per nanometer for each lumen as:

Photosynthesis - EQN 1

where Wrel(λ) is the relative spectral power distribution, V(λ) is the luminous efficiency function at wavelength λ, and Δλ is the wavelength interval (typically 5 nm). For the above example, the spectral radiant flux per nanometer for each lumen at 440 nm is 22.5 microwatts, while the total radiant flux per lumen is 3.18 milliwatts.

Photosynthesis - FIG 4

FIG. 5 – CIE 1931 luminous efficiency function V(λ)

With this, we can calculate the photosynthetic photon flux (PPF) per nanometer in micromoles per second per nanometer:

Photosynthesis - EQN 2

(where Na is Avogardo’s constant), while summing over the range of 400 nm to 700 nm yields the photosynthetic photon flux (PPF) per lumen for the given light source:

Photosynthesis - EQN 3

Given an illuminance value (lumens per square meter) and knowing the light source SPD, we can similarly calculate the photosynthetic photon flux density (PPFD) in micromoles per second per square meter (μmol/sec-m2) for the given light source. Again for the above example, one kilolux is equal to 14.62 μmol/sec-m2.

Conversion Factors

It is easy enough to find graphical representations of light source spectral power distributions, but it is considerably more difficult to find this information in tabular form suitable for the above calculations. Fortunately, this information is published in CIE 15:4, Colorimetry (CIE 2004). It does not include white light LEDs, but this information can be obtained by digitizing manufacturers’ product catalog data (e.g., Philips 2014a).

Given such information, it possible to calculate kilolux-to-PPFD conversion factors for common light sources:

Light Source Conversion Factor
CIE A (incandescent, 2856K) 20.3
CIE 5000K daylight (D50) 18.1
CIE 5500K daylight (D55) 18.1
CIE 6500K daylight (D65) 18.3
CIE 7500K daylight (D75) 18.6
CIE HP1 (standard high-pressure sodium, 1959K) 11.7
CIE HP2 (color-enhanced high-pressure sodium, 2506K) 19.3
CIE HP3 (high-pressure metal halide, 3144K) 14.4
CIE HP4 (high-pressure metal halide, 4002K) 15.0
CIE HP5 (high-pressure metal halide, 4039K) 16.3
2700K white light LED (Philips Luxeon Rebel LXW9-PW27) 18.1
3000K white light LED (Philips Luxeon Rebel LXW9-PW30) 17.1
3500K white light LED (Philips Luxeon Rebel LXW7-PW35) 14.6
4000K white light LED (Philips Luxeon Rebel LXW8-PW40) 14.3
5000K white light LED (Philips Luxeon Rebel LXW8-PW50) 14.6

Table 2 – Illuminance (kilolux) to PPFD (μmol/sec-m2) conversion factors

Table 2 does not include commercial products such as the Sylvania SHP-TS Grolux (with a CCT of 2050K) because Sylvania and most other lamp manufacturers do not publish their lamp SPDs in tabular form. It is possible to digitize the graphical representations of white light LEDs because the bandwidth of the blue “pump” LEDs is at least 15 nm. With high-pressure sodium and metal halide lamps, however, it is impossible to digitize their published SPDs because the wavelength resolution is unknown. A subnanometer-wide line emission, for example, could vary in height by five times, depending on whether the wavelength binning is 1 nm or 5 nm.

Overcast Skies

Table 2 presents conversion factors for the CIE Clear Sky with CCTs ranging from 5000K to 7500K. Their spectral power distributions (shown in the Appendix) were calculated in accordance with the equations published in CIE 15:4, Colorimetry (CIE 2004), which were in turn derived from the spectral distributions of 622 samples of daylight (skylight, and sunlight plus skylight), as discussed in Judd et al. (1964). Considering the variability of daylight, these SPDs are sufficient for most purposes.

What however about overcast skies? To answer this question, we reference “Colors of the Daytime Overcast Sky” by Lee and Hernández-Andréz (2006), who defined “overcast” as meeting two criteria: 1) no clear sky can be visible anywhere; and 2) cloud cover must be sufficiently optically thick that any cast shadows are indistinct.

The authors made over 9,100 spectral irradiance measurements in Granada, Spain, and Annapolis, Maryland on 40 overcast days, including days with drizzle, light rain and snow, with the Sun at least five degrees above the horizon. While the paper offers many interesting details, two items are of particular interest.

First, the correlated color temperature of overcast skies that the authors measured ranged from 5800K to 9300K, with their typical overcast skies having CCTs ranging from 6000K to 6600K. Second, the authors provided SPDs of their two most extreme skies, with CCTs of 5800K and 9300K. These SPDs were used to calculate illuminance (kilolux) to PPFD (μmol/sec-m2) conversion factors of 17.8 and 20.5 respectively. From these values, the CCTs of typical overcast skies can be linearly interpolated to be 18.2 ± 0.2. In other words, no different from typical clear skies.

LED Lighting for Horticulture

At this time, high-pressure sodium (HPS) lamps are the most common light source for greenhouse lighting, where it is commonly used to supplement daylight during the winter months. However, with the growing interest in urban horticulture that relies exclusively on electric lighting, light-emitting diodes offer many advantages. This is particularly true for multilayer cultivation, where the close spacing of plants in vertical rack-mounted trays make HPS lighting impractical.

McCree (1972a) noted that the relative quantum yield for crop plant photosynthesis has two peaks at 440 nm and 620 nm. He also noted however, the Emerson effect, which states that photosynthesis in the presence of two or more wavelengths can be more efficient than the sum of that due to the individual wavelengths. In particular, adding white or red light (less than 680 nm) to deep red light (greater than 680 nm) can beneficially increase the rate of photosynthesis.

Green light is also used in photosynthesis, as can be seen from the leaf action spectrum (FIG. 2). It has been established that green light drives photosynthesis more effectively than red or blue light deep within the leaf (Terashima et al. 2009). Further, the insects used in greenhouses as pollinators and biological control agents see best in the green and ultraviolet regions of the spectrum. More interestingly, changes of even 10 nm in the peak wavelength of green light can have dramatic effects on the growth of plants such as lettuce (Johkan et al. 2012).

Plants also exhibit photomorphological responses (i.e., growth and development) to ultraviolet radiation (Zuk-Golaszewska et al. 2003). In the past, this has been mostly of theoretical interest to botanists. Now however with the rapid commercialization of ultraviolet LEDs (Shih 2015) with wavelengths covering the plant biologically-active spectrum of 280 nm to 400 nm, UV-A and UV-B LEDs will likely also find application in horticultural lighting.

It is likely for this reason that many horticultural LED modules feature efficient 450 nm indium-gallium-nitride (InGaN) deep blue LEDs and 660 nm aluminum-indium-gallium phosphide (AlInGaP) deep red LEDs. Typical examples of these LEDs are the Philips Luxeon Royal Blue (LXML-PRO1-0425) and Deep Red (LXM3-PD01) products (Philips 2014b). Both of these products are quite efficacious, converting some 45% of their electrical input power into visible light. Green LEDs, while beneficial, are rarely used because of their much lower radiant efficacies. (This may soon change, however, as OSRAM Opto recently announced the development of 530 nm InGaN green LEDs with 25% external quantum efficiency.)

Herein however lies a problem: 450 nm and 660 nm are close to the limits of our color vision (see FIG. 4). Consequently, Philips and other manufacturers typically express the optical performance of these products in radiometric rather than photometric terms – milliwatts instead of lumens.

So, the lighting design process becomes a bit more complicated. We first need to digitize the published LED spectral power distributions to determine the conversion factors between milliwatts and lumens – these will be needed for the lighting design simulations. These are given by:

Photosynthesis - EQN 4

where ΦL is the luminous flux, ΦR(λ) is the relative spectral radiant flux and V(λ) is the luminous efficiency function at wavelength λ.

Using the Philips Luxeon Royal Blue and Deep Red products as an example, the respective conversion factors are approximately 0.07 and 0.03 lumens per milliwatt (lm/mW). However, these figures must be approached with some caution, as they apply to 450 nm and 660 LEDs only. If for example the peak wavelength of deep blue LED was 440 nm rather than 450 nm, the conversion factor would be 0.05 lm/mW. Similarly, if the peak wavelength of the deep red LED was 650 nm rather than 660 nm, the conversion factor would be 0.06 lm/mW. The Philips LED binning ranges are 440 to 460 nm and 650 to 670 nm respectively, which equates to (from FIG. 4) conversion factor uncertainties of +75%, -50% for blue and +60%, -30% for red. The above conversion factors are therefore decidedly approximate.

(Some horticultural LED module manufacturers bin their LEDs more tightly, as peak maxima shifts as small as 10 nm have been shown to have dramatic effects on plant growth. Unless however the binning policy is stated in the manufacturer’s product literature, this cannot be assumed.)

A further word of caution: even the best illuminance meters can be wildly inaccurate when measuring deep blue and deep red light levels. Commercially available photometers are usually classified according to their f1 number (with f1  < 3% being preferred), which is basically a measure of how closely the spectral response of the meter matches that of the photopic visual efficiency function (FIG. 4). As noted in CIE 127:2007, Measurement of LEDs (CIE 2007), this is useful for white light measurements only. To quote, “In the case of single-color LEDs, the spectral mismatch errors can be very large even if f1 is reasonably small, due to the fact that some LED spectra are peaking in the wings of the V(λ) function where the deviation makes little effects on f1 but can cause large errors.”

With these conversion factors in hand, we can now calculate the illuminance-to-PPFD conversion factors for horticultural LEDs:

Light Source Conversion Factor
450 nm deep blue LED 119.4
525 nm green LED 8.4
660 nm deep red LED 130.5

Table 3 – Illuminance (kilolux) to PPFD (μmol/sec-m2) conversion factors

How horticulturalists choose to balance the ratio of red to blue light will likely depend on the specific plant species being cultivated and their stage of growth. Some plants like shade, while others prefer direct sunlight, with different SPD requirements. In addition, far-red 735 nm LEDs may be employed to induce flowering. Regardless, the above conversion factors will still be useful.

In addition to using chlorophylls and carotenoids for photosynthesis, plants use these and other photopigments for a wide variety of functions. The phytochromes Pr and Pfr, for example, respond to 660 nm red and 735 nm infrared radiation respectively, and in doing so induce seed germination and flowering, regulate leaf expansion and stem elongation, and trigger photoperiod and shade avoidance responses (see Appendix A).

Other photopigments regulate phototropism (leaf and stem orientation) and circadian rhythms (for which blue light is the most effective), photomorphogenesis (plant shape), root growth, stomatal opening, chloroplast movement … the list goes on, as horticultural researchers continue to explore the role between lamp SPDs and optimal plant health and growth. (See www.photobiology.info for an informative summary of plant photobiology.)

Summary

As a reminder, photosynthetically active radiation (PAR) does not consider the spectral response of plants (FIG. 3); it simply represents the number of photons (quanta) per unit area per second within the range of 400 to 700 nm. With the availability of color-tunable LED modules for greenhouse lighting, horticulturalists will likely want to experiment with different SPDs for specific crops and flowering plants, as well as both the directionality and daily timing (photoperiods) of the luminaires. Regardless, being able to convert predicted and measured illuminance values to PPFD values for common light sources will certainly ease the communication problem between lighting designers and horticulturalists.

Acknowledgements

Thanks to Tessa Pocock of the Smart Lighting Engineering Research Center, Rensselaer Polytechnic Institute, for her review and comments on this article.

Appendix A – Photosynthesis and Visible Light

For illumination engineers, it might seem suspicious that the photosynthetically active radiation is defined over the spectral range 400 nm to 700 nm – exactly the range we commonly assume for human vision. What about longer and shorter wavelengths?

When McCree [4] measured his 22 crop species both in the field and in laboratory growth chambers, he obtained the following action spectra:

Photosynthesis - FIG A1

Figure A1 – Growth Chamber Action Spectra

Photosynthesis - FIG A2

Figure A2 – Field Action Spectra

which clearly explain the logic of the 400 – 700 nm spectral range.

Below 400 nm, there is the risk of photooxidation that generates toxic radicals, which can destroy the cell’s chlorophyll and other cellular components. Under intense UV radiation, violaxanthin (which is involved in photosynthesis) is converted via the xanthophyll cycle into zeaxanthin. In doing so, it receives excess energy from chlolorphyll and releases it as heat. This process thereby offers the plant photoprotection.

At the same time, other plant photopigments, including cryptochromes and phototropins, do have sensitivities (as measured in vitro) that extend into the ultraviolet, and likely respond under dim light conditions. However, these are likely suppressed under high light conditions by the xanthophyll process.

Above 700 nm, the photon energy is too low to activate the photosynthetic process via the chlorophylls and various cartenoids. However, the phytochrome photopigment, which is responsible for stem elongation, leaf expansion, shade avoidance, neighbor perception, seed germination, and flower induction, has two isoforms called Pr and Pfr. In its ground state Pr, phytochrome has a spectral absorbance peak of 660 nm. When it absorbs a red photon, it converts to its Pfr state, which has a spectral absorbance peak of 730 nm. When the phytochrome molecule absorbs a far-red photon, it converts back to its Pr state, and in doing so triggers a physiological change in the plant.

Photosynthesis - FIG A3

Figure A3 – Phytochrome Action Spectra

Chlorophyllous leaves are transparent to infrared radiation, are so the phytochrome signaling mechanism is ideal for sensing the lighting environment on forest floors and in the presence of neighboring plants competing for available direct sunlight.

Appendix B – Light Source SPDs

Photosynthesis - CIE A SPDPhotosynthesis - CIE D50 SPDPhotosynthesis - CIE D55 SPDPhotosynthesis - CIE D65 SPDPhotosynthesis - CIE D75 SPD Photosynthesis - 2700K LED Photosynthesis - 3000K LED Photosynthesis - 3500K LED Photosynthesis - 4000K LEDPhotosynthesis - 5000K LEDPhotosynthesis - CIE HP1 SPD Photosynthesis - CIE HP2 SPD Photosynthesis - CIE HP3 SPD Photosynthesis - CIE HP4 SPDPhotosynthesis - CIE HP5 SPDPhotosynthesis - Horticultural LEDs

References

Barnes, C., T. Tibbitts, J. Sager, G. Deitzer, D. Bubenheim, G. Koerner, and B. Bugbee. 1993. “Accuracy of Quantum Sensors Measuring Yield Photon Flux and Photosynthetic Photon Flux,” Horticultural Science 28(122):1197-1200.

Biggs, W. W., A. R. Edison, Jerry D. Eastin, K. W. Brown, J. W. Maranville, and M. D. Clegg 1971. “Photosynthesis Light Sensor and Meter,” Ecology 52:125–131.

CIE. 2004. Colorimetry, Third Edition. CIE Technical Report 15:2004. Vienna, Austria: Commission Internationale de l’Eclairage.

CIE. 2007. Measurement of LEDs, Second Edition. CIE Technical Report 127:2007. Vienna, Austria: Commission Internationale de l’Eclairage.

Johkan, M, K. Shoji, F. Goto, S. Hahida, and T. Yoshihara. 2012. “Effect of Green Light Wavelength and Intensity on Photomorphogenesis and Photosynthesis in Lactuca sativa,” Environmental and Experimental Botany 75:128-133.

Judd., D. B., D. L. MacAdam, and G. Wyszecki. 1964. “Spectral Distribution of Typical Daylight as a Function of Correlated Color Temperature,” Journal of the Optical Society of America 54(8):1031-1039.

Lee, R. L., and J. Hernández-Andrés. 2006. “Colors of the Daytime Overcast Sky,” Applied Optics 44(27):5712-5722

Massa, G. D., H.-H. Kim, R. M. Wheeler, and C. A. Mitchell. 2008. “Plant Productivity in Response to LED Lighting,” Horticultural Science 43(7):1951-1956.

McCree, K. J. 1972a. “The Action Spectrum, Absorptance and Quantum Yield of Photosynthesis in Crop Plants,” Agricultural and Forest Meteorology 9:191-216.

McCree, K. J. 1972b. “Tests of Current Definitions of Photosynthetically Active Radiation Against Leaf Photosynthesis Data,” Agricultural and Forest Meteorology 10:443-453.

Mitchell, C. A., A.-J. Both, C. M. Bourget, J. F. Burr, C. Kubota, R. G. Lopez, R. C. Morrow, and E. S. Runkle. 2012. “LEDs: The Future of Greenhouse Lighting!”, Chronica Horticulturae 52(1):6-11.

Nelson, J. A., and B. Bugbee. 2014. “Economic Analysis of Greenhouse Lighting: Light Emitting Diodes vs. High Intensity Discharge Fixtures,” PLOS One 9(6).

Philips Lumileds. 2014a. Luxeon Rebel ES Portfolio Datasheet DS61. Philips Lumileds Lighting Company.

Philips Lumileds. 2014b. Luxeon Rebel Color Portfolio Datasheet DS68. Philips Lumileds Lighting Company.

Shih, M. 2015. “Emerging Applications for UV LEDs Drive Broad Interest,” LEDs Magazine 58(11/12):19-22 (Nov/Dec).

Terashima, I., T. Fujita, T. Inoue, W. S. Chow, and R. Oguchi. 1009. “Green Light Drives Leaf Photosynthesis More Efficiently than Red Light in Strong White Light: Revisiting the Enigmatic Question of Why Leaves are Green,” Plant & Cell Physiology 50(4):684-697.

Thimijan, R. W., and R. D. Heins. 1983. “Photometric, Radiometric, and Quantum Light Units of Measure: A Review of Procedures for Interconversion,” Horticultural Science 18(6):818-822.

Zhu, X.-G., S. P. Long, and D. R. Ort. 2010. “Improving Photosynthetic Efficiency for Greater Yield,” Annual Review of Plant Biology 61:235-261.

Zuk-Golaszewska, K., M. K. Upadhyaya, and J. Golaszewski. 2003. “The Effect of UV-B radiation on Plant Growth and Development,” Plant Soil Environ 49:135-140.

[1] A mole is a unit of measurement used in chemistry to express the number of elementary entities in a substance that is equal to the number of atoms in  12 grams of the isotope carbon-12. It corresponds to the Avogadro constant, whose value Na is 6.022 x 1023 particles (in this case photons) per mole. A micromole is one millionth of a mole. (A micromole [μmol] of photons was sometimes referred to by plant scientists as a microeinstein. However, this unit of measurement is not part of the International System of Units (SI), and so its use has been deprecated.)

[2] The quantum yield in photosynthesis is defined as the micromoles of carbon dioxide fixed per micromole of photons absorbed.