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anA QoQ =
0.90
0.80
0.70
0.60
0.50
0.40
Relative Quantum Yield
0.30 —#— Photon-weighted
0.20 ~Energy-Weighted
0.10
0.00
Photometry and Photosynthesis
by Ian Ashdown | 2019 Feb Fri | Horticulture
Tan Ashdown, P. Eng., FIES, Senior Scientist, SunTracker Technologies Ltd.
Published: 2014/12/10
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 lux-to-PPFD conversion factors for
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orticultural 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 (IIPS) 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 fi light fi
into chemical energy that is used for growth and development. All that is needed for this
process is carbon dioxide (CO.), 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 ct 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
proc:
ss of photosynthesis to fix the carbon in CO, 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
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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.
100.0 7
900
80.0
ZF 700
3 600 —Chlorophyll A
= soo —Chlorophyll B
a
£ 400 Beta-carotene
a
2 300 Phytochrome (Pr)
200 —Phytochrome (Pfr)
10.0
0.0
300 400, 500 600 700 800
Wavelength (nm)
FIG. 1 - Photopigment spectral absorptanees.
MeCree (1972a) measured the spectral absorptance (FIG. 2) and quantum yield of Cop
assimilation for the leaves of 22 species of crop plants[2). Taking the average measurements
at 25 nm intervals forall plant species (Table 1), he produced the photon-weighted relative
quantum yield (Table 1) that is representative of most crop plants.
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pea
2°
°
°
Absorptance
e oo
GS
°
2
°
400 450 500 550 600 650 700
‘Wavelength (nm)
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
525 072
550 0.82
575 o.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 A is given by the Planck-Einstein relation:
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‘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, itis
synonymous with irradiance.) As shown by McCree (1972), photon-weighted PPED 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.
1.00
0.90
0.80
0.70
0.60
0.50
0.40
Relative Quantum Yield
0.30 —— Photon-weighted
0.20 ~Energy-Weighted |__|
0.10
0.00
400 450 500 550 600 650 700
Wavelength (nm)
FIG. 3 ~ Relative quantum yield for erop 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
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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).
1.000
0.900
0.800
0.700
0.600
0.500
0.400
0.300 VV
0.200 +++}
Relative SPD
0.100
0.000 +41 jj ttt
400 450 500 550 600 650 700
Wavelength (nm)
FIG. 4 ~ 500K 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 funetion (FIG. 5), we can calculate the spectral radiant flux (2) in
watts per nanometer for each lumen as:
Wrei(A)
683 + 7H V AWrei(AA
@(A)/Im =
where Wye{(A) is the relative spectral power distribution, VW) is the luminous efficiency
function at wavelength A, and Ad is the wavelength interval (typically § 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.
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oa
400 450 500 550 600 650 700
Wavelength (nm)
FIG. 5 ~ CIE 1931 luminous efficiency function VO)
With this, we can calculate the photosynthetic photon flux (PPF) per nanometer in
micromoles per second per nanometer:
Ao (A)
=10-%*
PPR /nm = 10~° «yap
(where Ni 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:
700 700
10%
PPF= A@(A)AA = 8.359 * 10 « > AM(ADAA
Ware" Dy 2
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 (timol/sec-m_) for the given light source. Again for the above
example, one lux is equal to 0.01462 umol/sec-m’.
Conversion Factors
Itis easy enough to find graphical representations of light source spectral power
listributions, but it is considerably more difficult to find this information in tabular form
suitable for the above calculations. Fortunately, this information is published in CTE 15:4,
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Conversion
Light Source Factor
CIE A (incandescent, 2856K) 0.0203
CIE 5000K daylight (D50) 0.0181
CIE 550K daylight (D55) 0.0181
CIE 6500K daylight (D65) 0.0183,
CIE 7500K daylight (D75) 0.0186
CIE HP! (standard high-pressure sodium, 1959K) 0.on7
CIE HP2 (color-enhanced high-pressure sodium, 2506K) 0.0193,
CIE HP3 (high-pressure metal halide, 3144K) 0.0144
CIE HP4 (high-pressure metal halide, 4002K) 0.0150
CIE HP5 (high-pressure metal halide, 4039K) 0.0163
2700K white light LED (Philips Luxeon Rebel LXW9-PW27) 0.0181
3000K white light LED (Philips Luxeon Rebel LXW9-PW30) oor
3500K white light LED (Philips Luxeon Rebel LXW7-PW35) 0.0146
4000K white light LED (Philips Luxeon Rebel LXW8-PW40) 0.0143,
5000K white light LED (Philips Luxeon Rebel LXW8-PW50) 0.0146
‘Table 2 - luminance (lux) to PPFD (umol/sec-m’ ) 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, its 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 sutfi
it for most purposes.
What however about overcast skies? To answer this question, we reference “Colors of the
Ravtime Qvernact Sle
* hy Tae and Harninder-Andrée (onnk) whe defined “avareact” ae
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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 900K, with their typical overcast skies having CCT’s ranging from 6000K to
6600K. Second, the authors provided SPDs of their two most extreme skies, with CCTs of
5800K and 900K. These SPDs were used to calculate illuminance (lux) to PPED (tumol/sec~
2
m_) conversion factors of 0.0178 and 0.0205 respectively. From these values, the CCTs of
typical overe:
tt skies can be linearly interpolated to be 0.0182 + 0.0002. 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 ean be more efficient than the
sum of that duc 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 ean b
2). Ithas 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
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Royal Blue (LXML-PRO1-0425) and Deep Red (LXM3-PDo1) 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:
700
, = 0.683 + win PaAVA)
1 = 0.683 + A
Lioo¥A)
where is the luminous flux, & g() is the relative spectral radiant flux and VQ) is the
luminous efficiency function at wavelength A.
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 (Im/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 Im/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
Im/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
é, -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 f, number (with f, < 3% being preferred), which is
basically a measure of how closely the spectral response of the meter matches that of the
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With these conversion factors in hand, we can now calculate the illuminance-to-PPED
conversion factors for horticultural LEDs:
Light Source Conversion Factor
450 nm deep blue LED o.01194
525 nm green LED 0.00084
660 nm deep red LED 0.01305
2
‘Table 3 — Iluminance (lux) to PPD (umol/sec-m-) 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 Py and Pf, 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
thythms (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
ssponse 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
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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:
350 400 450 S00 550. 600.650.700.750
Wavelength (nm)
Figure At — Growth Chamber Action Spectra.
1.00
0.90
0.80
0.70
0.60
0.50
0.40
0.30
0.20
0.10
0.00
Relative Action (a.u.)
350 400 450 500 550 600 «650. 700-750
Wavelength (nm)
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 cellis chlorophyll and other cellular components. Under intense UV radiation,
violaxanthin (which is involved in photosynthesis) is converted via the xanthophyll cycle into
zeaxanthin, Tn doing so, it receives excess energy (rout chlolorphyll and releases it as heat.
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‘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 eartenoids. 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 P, and Pf,. In its ground state Py,
phytochrome has a spectral absorbance peak of 660 nm. When it absorbs a red photon, it
converts to its Pf, state, which has a spectral absorbance peak of 730 nm. When the
phytochrome molecule absorbs a far-red photon, it converts back to its P, state, and in doing
so triggers a physiological change in the plant.
100.0
90.0
80.0
70.0
60.0
50.0
40.0
30.0
20.0
10.0
0.0
— Phytochrome (Pr)
~ ~~~ Phytochrome (Pf)
ive Absorptance (a.u.)
600 650 700 750 800
Wavelength (nm)
Figure Ag — 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
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CIEA
1.00
80
080
g 060 |
g 050
2 040
020
a0
00 | |
400 450 500 550 500 650 700
‘Wavelength (nm)
CIE DSO
140.00
120.00
Reative SPD
40.00
20.00
0.00
400 450 500 550 600 650 700
Wavelength (am)
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CIE DSS.
140.00
120.00
100.00
Relative SPD
g
3
20.00 | |
a |_| |
400 450 500 550 600 650 700
Wavelength (nm)
80.00 |
40.00
CIE D65
1140.00
120.00
10000 |.
20.00
60.00
Relative SPD
40.00
20.00
0.00
400 450 500 550 600 650 700
Wavelength (nm)
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CIE D75
140.00
120.00
300.00
60.00
Relative SPD
40.00
20.00
0.00
400 450 500 550 600 650 700
Wavelength (am)
2700K LED
1.00
090
080
070
goo
£050 }
2 040
030
020
0.00 ———
400 450 500 550 600 650 700
Wavelength (nm)
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3000K LED
1.00
0.30
0.80
070
0.60
050
Relative SPO
0.40
0.30
0.20
020
0.00
400 450 500 550 600 650 700
Wavelength (nm)
3500K LED
41.00
090
080
070
goo
£050
2 oao
030
020
0.0
0.00
400 450 500 550 600 650 700
Wavelength (nm)
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100
oso
oo
a0
060
$os0
i
3 ovo
030
020
oxo
0.00
1.00
030
80
070
030
020
010
0.00
‘You searched for photometry and pholosynthas's - All Things Lighting Assocation
4000K LED
400 450 00 550 600 650 700
Wavelength (nm)
5000K LED
400 450 500 350 600 650 700
‘Wavelength (nm)
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CIE HP1
350.00
Relative SPD
400 450 500 550 600 650 700
Wavelength (nm)
CIE HP2
Relative SPD
400 450 500 550 600 650 700
Wavelength (om)
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Relative SPD
Relative SPD
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CIE HP3
350.00
300.00
250.00
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15000 |
100.00
50.00
0.00
400 450 500 550
Wavelength (nm)
650 700
CIE HP4
vas |
= |
- |
50.00
0.09
450 500 350
Wavelength (am)
600 650 700
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CIE HPS.
350.00
8
i
z
250.00 |
5000
0.00
400 450 500 350 600 650 700
Wavelength (nm)
Horticultural LEDs
oso |
——450 nm lve
0.80
— =525 nmGreen
070
= = 660 nm Red
0.60
oso
40
600 700
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
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CIE. 2007. Measurement of LEDs, Second Edition. CIE Technical Report 127:2007. Vienna,
Austria: Commission Internationale de l’Eclairage.
Johkan, M, K. Shoji,
Wavelength and Intensity on Photomorphogenesis and Photosynthesis in Lactuea sativa,”
P. Goto, 8. Hahida, and T. Yoshihara. 2012. “Effect of Green Light
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. Herndndez-Andrés, 2006. “Colors of the Daytime Overcast Sky,” Applied
Opties 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, 8. Runkle, 2012. “LED:
52(1):6-11,
the Future of Greenhouse Lighting!” Chronica Horticulturae
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 D861. 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.
Photos
Chow, and R. Oguchi. 1009. “Green Light Drives Leaf
nthesis 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
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‘Zuk-Golaszewska, K, M. K. Upadhyaya, and J. Golaszewski. 2009. “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 Ng is 6.022 x 10°?
particles (in this case photons) per mole. A micromole is one millionth of a mole. (A
jicromole [zmol] 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 (SD, 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.
Botanical Light Pollution
by Ian Ashdown | 2019 Feb Wed | Horticulture
Tan Ashdown, P. Ens
FIES, Senior Scientist, SunTracker Technologies Ltd.
Published: 2016/02/17
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 A
factors, even low levels of light trespass from roadway and outdoor area luminaires can have
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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 — ean reduce erop yield by 20 to
{40 percent due to delayed flowering and ripening (Chen et al. 2009).
FIG. 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
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‘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
+ Dormaney
‘Taken together, these functions basically outline the life eycle from seed to adult plant.
‘The sum of these
jht-induced changes is called photomorphogenesis. There are other
photopigments involved, including blue light-sensitive cryptochromes (Lin 2002) and
ultraviolet-sensitive UVRS 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 Py
and Pf (e.g., Smith 2000). The Py 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 Pf; 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 Pj; isoform absorbs a far-red photon, it reverts to its P, isoform. Thus,
phytochrome performs the function of a resettable biological switch to initiate or terminate
photomorphological processes.
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uu
0.90
0.30
g 070
§ 0.60
Eoso
§ 0.20
030
0.20
0.10
0.00
Phytochrome (Pr)
Abs
— —Phytochrome (Pfr)
Wavelength (nm)
FIG. 2 ~ Phytochrome absorption spectra. (Source: Plants in Action, First Edition).
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 phy
iological reaction to the length of the day. Like humans and all other animals,
plants have cireadian 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
Pj to revert to Py, 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 P,. 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 ean
bea 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
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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 (c.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
botanical light pollution. For
environment is not a good thing. We can call it what it
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), 50 it is reasonable to ask whether
this influences (or “screens”) the phytochrome absorptance spectra in vivo.
100.0
90.0
80.0
70.0
60.0 —Chlorophyll A
50.0 — Chlorophyll B
40.0 ——~ Beta-carotene
Absorptance (a.u.)
30.0 — Phytochrome (Pr)
ana fm
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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
emarkable correlation with
their results as induction and reversion “action spectra” shows
the in vitro absorptance spectra of phytochrome (FIG. 4).
3.8 8
we. quantum responsivity
s
10!
350 400 500 600 700 ‘800
Woveiength [nm]
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,
Fora light source, we typically have its relative spectral power distribution (SPD), which is,
measured in watts per nanometer. However, from the Planck-Hinstein 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)
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| .ev
030
080
070
0.60
oso
040
Radiant Flux
— — Photon Flux
030
020
010
0.00
400 450 S00 550 600 650 700
Wavelength (nm)
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:
1, Multiply the SPD values of each lamp by the CIE 1931 luminous efficiency funetion VO)
shown in FIG. 6 from 400 nm to 700 nm.
2, Sum the results of Step 1 to obtain the relative lumens Prof and test generated by the
two lamps.
3. Multiply the SPD values of the test lamp by ®rof / Ptest-
‘The two SPDs now represent the same number of photopic lumens (ie., 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 P; spectral absorptance spectrum.
3. Sum the results of Step 5 to obtain the P, action values pAref,rand PAtest,r-
4. Multiply the results of Step 4 by the phytochrome Pf, spectral absorptance spectrum.
5.Sum the results of Step 7 to obtain the Pf action values PAret fr and PAtest fr
and finally:
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1.00
0.90 |
30.80
3070
@ 0.60 i
2050 |
Fao |
20.30 \
$020
0.10
0.00
400 450 500 550 600 650 700
Wavelength (nm)
FIG. 6 ~ CIE 1931 lun
1ous efficiency function V(_).
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) itis 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 Pf; is only 60 percent that of P,, 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 nawaat than tha Fallen
See rata i leeireaesarsara
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000K 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 sourees.
‘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.
a
ogo |__|
oso
—noxito
—sco.t0
—ssc0c.t0
i a
ast scant .t0
as
ooo
“oo 0 exo 700 so
Wavelength (nm)
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. Pre PiaPy
High-pressure sodium | 1.0 1.0 1.0
‘2700K white light LED | 1.7 23 19
3000K white light LED | 1.5 2.0 17
3500K white light LED | 1.0 12 41
1.0 10 [1.0
‘5000K white light LED | 0.9 10 09
‘Table 2 — Relative phytochrome action values.
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It should be noted however that these results apply to Lumileds LUXEON products only.
Looking at FIG. 7, itis 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 Ro 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 000K. 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.
1.00
o90
200k LED
— —Whiessed
200 soo 600 700 300
‘Wavelength (am)
FIG. 8 — 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-
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Light Source | Relative Phyt
P._
[WhitetredLED [09 09 [09
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 000K 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
chlorophyll A.)
for spectral overlapping between phytochrome P, and
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 P, 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).
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1.00
030
0.80
070
0.60
050
White Seedlings
Relative Action
2
é
— —Green Seedlings
0.30
0.20
0.10
0.00
400 500 600 700
Wavelength (nm)
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.
. artificial) red radiation will convert the P isoform in the exposed plant to
Any excess (
fy , while any excess far-red radiation will convert the Pf, isoform to P;, Either aetion will
upset the plantis 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
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