How Plants Use Light
Human and plant light perception use many of the same
molecules, however, our eyes are more easily fooled. Narrow band red, blue, and
green light when mixed in the right proportion are perceived as white light to
the human eye. However, a plant is quite aware of the fact that it is receiving
three individual spectra and its growth habit will demonstrate that perception.
Broad spectrum white light can come in many forms depending on the source. To
the human eye, we mostly perceive these different spectra as cooler (blue) or
warmer (orange/red) depending if it is a metal halide or high-pressure sodium
lamp, or in the case of LEDs and fluorescent bulbs, what type of phosphor
coating is used. To a plant however, each individual wavelength may promote a
different growth habit and photomorphogenic response.
Incoming photons are absorbed by pigments, which absorb light
as energy, and photoreceptors which perceive light as a signal. When absorbed
by the most known pigment, chlorophyll, photons can be used to drive
photosynthesis and growth. However, chlorophyll alone with its wide-ranging
absorption spectrum is not enough to efficiently harvest light. The “antenna
complex” is a concept that describes how accessory pigments such as carotenoids
can assist both in capturing light that chlorophyll does not absorb, or
dissipating excess light as heat (non-photochemical quenching) when
photosynthetic reaction centers are overloaded with incoming energy.
Accessory pigments are primarily carotenoids such as
beta-carotene, lutein, zeaxanthin, antheraxanthin, and violaxanthin. These
pigments are yellow to orange in color and absorb most strongly in the range of
450 nm to 550 nm. Some of these pigments change forms based on lighting
conditions through processes called epoxidation and de-epoxidation. If fluence
is too high, damage can occur to the photosynthetic apparatus, so it is
important for a plant to be able to deal with this incoming energy. Under pure
sunlight where fluence may fluctuate throughout the day, the antenna complex
adjusts to accept or dissipate light. When fluence is low, violaxanthin will
capture photons and transfer this energy to chlorophyll, improving efficiency
of light absorption. When fluence is high, violaxanthin is de-epoxidated (converted)
into zeaxanthin which then dissipates excess photons as heat. Beta-carotene
functions similarly to violaxanthin and lutein functions like zeaxanthin but
without this interconversion process called the “Xanthophyll Cycle.” This flow
of energy between pigments occurs spontaneously as they become “excited” by
photons. Interestingly, carotenoids that protect plants from light and improve
their ability to capture light can also serve similar functions within the eyes
of many animal species. There are several other plant pigments unassociated
with the photosynthetic light-harvesting complex including anthocyanin and
lycopene. Though these compounds do absorb light, their main function is to
protect cells and DNA from damaging UV radiation as well as scavenge “free
radicals” such as hydrogen peroxide, preventing further cellular damage.
Photoreceptors in most cases are proteins paired with a
“chromophore” that absorbs certain wavelengths of light and then sends a signal
to the plant that influences photomorphogenesis. There are several different
types of photoreceptors and their light absorption ranges
overlap. Cryptochromes use light in the range of 300 nm to 500 nm
though it most strongly absorbs at 350 nm (UV-A) and 450 nm (blue). This receptor,
when excited by light, prevents elongation of hypocotyls (main stem of
seedlings) and even mediates flowering and photoperiod in some
species. Phototropins are also blue/UV-A absorbing photoreceptors but
with a much stronger absorption peak at 450 nm and are thought to regulate
phototropism (process in which plants move in response to light), stomatal
aperture (opening and closing), movement of chloroplasts (photosynthetic
organelles) within leaf cells, and inhibition of leaf
expansion. Phytochromes are some of the more famous photoreceptors as
they can strongly influence flowering. It is a little known fact that
phytochromes actually absorb light in the range of 300 nm to 800 nm. Most of
the known functions however, are a result of the absorption peaks at 660 nm in
the Pr form and 730 nm in the Pfr form. Phytochromes are constantly
changing form and reach a “photoequilibrium” (more information available in the
“Guide to Photomorphogenesis”) that is regulated by spectral ratio and PPFD
present in the growing environment. Depending on the photoequilibrium of
phytochrome, different signals are sent to metabolic pathways within the plant
that regulate many processes including germination, seedling establishment,
stem elongation, leaf expansion, and of course flowering and photoperiod.
Different ratios of R:FR light received by a plant will dictate how the plant
develops in terms of compactness, flower size, flower number, etc. There are
several other newly discovered and under-researched photoreceptors (UV-B
receptor correlated with anthocyanin accumulation) but these will not be
discussed in this article.
Since there is much overlap in the absorption spectra of
these photoreceptors, most photomorphogenic responses are co-regulated. Some
responses may be turned on and off by one receptor, but the expression of that
response can be amplified by another receptor. The enigmatic “circadian clock”
that regulates so many functions within plants is a culmination of activity
from multiple photoreceptors entraining a rhythm of growth patterns based on
photoperiod, light spectrum, and PPFD. This rhythm of growth patterns within a
plant strongly influences photomorphogenic outcomes however, just like
photosynthesis, there is an action spectrum for all photomorphogenic responses
that is dictated by a mixture of signals from these photoreceptors and does not
necessarily mirror the absorption spectrum.
Supplementing Specific Wavelengths Vs Broad Spectrum
When considering using narrow band lighting, the most
important things to consider are whether or not your plants are already exposed
to broad spectrum light (solar for greenhouses, or a broad-spectrum fixture for
sole-source lighting applications) and which crops you are growing. When plants
are already exposed to broad spectrum lighting from a sole-source fixture, it
makes sense to supplement with narrow band lighting only if there is a desired
photomorphogenic effect that your crop can’t achieve without being exposed to a
specific waveband. However, if you are growing under solar radiation and a high
DLI, your crop may not be as responsive to changes in light spectrum, as solar
radiation is quite broad already and may drown out the photomorphogenic
benefits of narrow band lighting. Another aspect to consider when growing under
solar radiation is whether or not you need to increase your DLI. If you
supplement with a narrow band fixture as a method of increasing DLI, you may
see inconsistencies in product quality as solar radiation increases and
decreases throughout the year exposing your crops to differing amounts of
sunlight and narrow band light. If your DLI is constant and you only wish to
induce a photomorphogenic response such as coloring, compactness, or rooting,
it may make sense for you to supplement more blue light. However, DLI often has
more of an impact on desired traits than spectrum. If you are growing a
flowering/fruiting crop and only wish to encourage more flower/fruit growth
(with a sufficient DLI and photoperiod) it may be beneficial to supplement more
red light, as 660 nm light encourages phytochrome responses in many species,
which sends signals throughout the plant to encourage reproductive growth.
Conclusion
Narrow band lighting can provide acceptable growth for many
species, so long as there are no fluctuations in DLI independent of the portion
supplied by the light fixture. However, plants use several different
photoreceptors and pigments that cooperatively regulate growth and development.
Plants developed these photomorphogenic responses under broad spectrum light
and it is very rare for a certain species to express a response to narrow band
lighting that cannot also be achieved by broad spectrum lighting given
sufficient DLI. For consistent product quality and capability to produce a
broad array of crop species without complications due to lighting, broad spectrum
fixtures are a safer choice. Different species can have varying responses to
changes in light spectrum. Research is constantly underway that helps us
understand how individual crops respond to different light spectra, and in some
cases there is clear evidence about what type of lighting is best for a crop.
If you are uncertain about the response your crop may have, supplementing with
broad spectrum light has a proven record of improving crop quality,
consistency, and yield.