Ok, admittedly I've only skimmed this thread, but I'll toss out some thoughts, for what they're worth. Apologies for the length too. This Wikipedia article should be helpful in understanding and visualizing the processes I’m describing:
http://en.wikipedia.org/wiki/Photosynthesis
Step 1 in photosynthesis is to capture photons. When a photon is absorbed it excites an electron from a low energy level to a higher energy level. It takes a particular quanta of energy to do this (though there is a bit of “wiggle room†in the wavelength that can be absorbed due to electron vibration), which is why specific photopigments only absorb around specific wavelengths. Once the electron is excited the energy is passed from electron to electron (through the antenna proteins) and to the reaction center of photosystem II (PSII). The reaction center includes a molecule of chlorophyll a and numerous proteins.
So step 2 of photosynthesis is transferring the energy from absorbed photons down antenna proteins to the reaction center of PSII.
Step 3 of photosynthesis is using that absorbed energy to do some work. An electron from the chl a in the reaction center is excited and transferred to a carrier protein. That leaves the reaction center oxidized (= lacking an electron). The same process happens 3 more times for a loss of 4 electrons total. Again, those electrons get transferred from the reaction center to a carrier protein (e.g., plastoquinone). Having lost 4 e- the reaction center needs to get back some electrons. To do that it splits 2 water molecules as in the reaction below:
2H2O = 4e- + O2 + 4H+
Doing so produces the 4 needed e-, reducing PSII. It also produces molecular oxygen and 4 protons. The protons are used later to produce ATP and the ATP is used to produce carbohydrates and do other cellular work.
To this point we still haven’t fixed any carbon, and we’re about half way toward having accomplished photosynthesis.
Step 4 of photosynthesis is transporting the electrons we generated above down an electron transport chain to photosystem I (PSI). At PSI similar processes of photon absorption, energy transfer, electron excitation and charge separation take place. Electrons are transferred from PSI to NADP. They are replaced by e- coming down the electron transport chain to PSI. NADP is reduced at PSI to NADPH by ferredoxin using the e- from PSI and an H+. NADPH is used as a reducing agent in various cellular activities, including carbon fixation.
Step 5 of photosynthesis is using the ATP generated above (H+ produced at PSII used to generate ATP) and the NADPH generated above (produced at PSI) to take CO2 and turn it into carbohydrates using the Calvin cycle.
Summarizing, for photosynthesis to happen we need 5 major steps to take place:
1) Absorb photons in antenna proteins of PSII.
2) Transfer the energy from the photons to PSII, use that energy to electrons from PSII to a carrier protein.
3) Replace those electrons by slitting water, and use the H+ that results to make ATP.
4) Transfer electrons from PSII down the electron transport chain to PSI. At PSI transfer electrons to NADP with ferrodoxin and H+ to make NADPH.
5) Use the ATP from PSII and the NADPH from PSI to fuel the Calvin cycle, turning CO2 into carbohydrates.
There are several conditions that can limit the rate of photosynthesis for a given organism at a given temperature:
1) The supply of photons to drive the whole process.
2) The supply of carrier proteins (plastoquinone) at PSII to take up and transfer electrons down the electron transport chain.
3) The supply of CO2.
The rate that photons are captured by PSII depends on two things:
a) the number of photons available (in other words, PAR/PPFD)
b) wavelength-specific absorption of those photons by photosynthetic pigments/accessory pigments
As you can see in the action spectrum on the previous page, the zooxanthellae in corals are best at absorbing "blue" photons, good at absorbing "violet", "cyan", and "red-orange" and aren't as good at absorbing "yellow" or "green" photons.
The particular action spectra depicted above is a bit more "spikey" than I would say is common for most corals. In other words, the difference in the likelihood of absorbing a blue vs. a green vs. a red photon in most zooxanthellae is somewhat small--smaller than depicted above. Of course, it depends on the suite of pigments the zooxanthellae are producing. Different zoox. in different situations, or the same zoox. in different situations, will have slightly different action spectra. As a side-note: zooxanthellae look brown because they are reflecting a mix of wavelengths heavy in the green, yellow and orange. Green plants have much “spikier†action spectra than dinoflagellates: they absorb heavily in the blue, violet, and red, and reflect heavily in the green and yellow. Hence, green plants look green and zooxanthellae look brown.
If we use the action spectrum above as a guide, the coral depicted above is about 4x as likely to absorb a blue photon as a yellow photon. It is about 3x as likely to absorb a red photon as a yellow photon, and about 3/4x as likely to absorb a red photon as a blue photon.
A
critically important point to consider that I’ll make here is that absorbing photons IS NOT equivalent to performing photosynthesis. It is merely the first step.
Let’s compare what happens at high levels to what happens at low light levels. With high light levels most of the reaction centers of PSII in a cell will have absorbed a photon and will have an excited electron ready to be transferred down the electron transport chain. At low light levels many of the reaction centers will not have absorbed a photon (simply because there are fewer photons available to absorb) and fewer will be ready to transport an electron.
The amount of time it takes for a photon to be absorbed by antenna proteins and transferred to PSII is very, very short (~10^-12 seconds). The amount of time it takes to transport an electron down the e- transport chain is still very short, but comparatively much longer (~10^-9 seconds)â€"about 1000x as long.
When the energy from a photon is transferred to PSII and an electron is excited, the electron only remains in that excited state for a short period of time (~10^-9 sec or less). If the electron is transferred to plastoquinone during that time it can be used in photosynthesis. However, if a plastoquinone molecule doesn’t get there in time then the electron falls back to its ground state. The energy from the absorbed photon is lost as heat and/or fluorescence and thereby doesn’t get used for photosynthesis.
When light levels are low there is lots of plastoquinone relative to the number of reaction centers that have excited electrons, so most of the energy can be put into photosynthesis. However, as more and more reaction centers absorb photons the plastoquinone pool starts missing more and more of the excited electrons before they return to the ground state. Eventually we reach a point where the electron transport chain is transferring electrons as fast as it possibly can. If we increase the light intensity we will continue to see more light absorbed, but that light energy is simply lost as fluorescence and heat and not put into photochemistry.
It takes a particular quanta of energy to excite and electron in PSII so that it can be transferred down the e- transport chain. If a photon has less energy than is required it can’t be used in photochemistry at all (e.g., less than ~700 nm). If, however, the photon yields more energy than is required to excite the electron in PSII the extra energy is lost from PSII as heat and fluorescence. Once a photon is absorbed it is doesn’t matter one bit what the wavelength was. A blue photon a green photon and a red photon can all excite exactly the same number of photons in PSII: one electron.
If the light intensity is high then the rate of photosynthesis will be limited by something such as the rate that electrons can be transported down the e- transport chain, not by a lack of light. If we have high light intensity of blue light then the zoox. will absorb more of the photon than they will with high intensity green light. However, since photosynthesis is not light-limited in this case, it doesn’t affect the rate of photosynthesis one way or the other (although, we could start to see photoinhibition if FAR more photons are absorbed than can be used).
If we are at or above the point of photosaturation, the spectrum of the light is therefore entirely irrelevant. If the organisms absorb more light, they will just dump more energy as heat and fluorescence. If they absorb less light they will dump less energy. They simply have no way to do anything with this additional energy.
Now, if we consider what happens at low light levels, then we have a somewhat different situation. Here we are light-limited. If more light is absorbed it WILL result in increased photosynthesis. Hence, at low light levels blue light may indeed yield higher rates of photosynthesis than green light. Practically speaking, given typical action spectra in zooxanthellae, the difference is so small it has essentially negligible effects considering what we as aquarists are trying to do (namely grow corals and such).
How to translate this into actually putting a light over an aquarium… Well, at higher light intensity, spectrum is irrelevant considering the spectrum of any bulb we might consider using. If the corals aren’t light limited, it simply doesn’t matter if they are absorbing lots and lots of blue photons, or a mix of fewer blue, violet, green, and red photons. They are getting as much light as they can use, and that’s that.
At low light levels spectrum may indeed matter, but not whole lot given the spectra of bulbs we use and typical action spectra for zooxanthellae.
If, however, we consider different spectra at DIFFERENT intensities (e.g., a bright 10 K metal halide vs. a dimmer 15 K halide…dimmer = lower PAR/PPFD) then we have a more complicated situation. To determine which bulb would produce higher rates of photosynthesis at potentially limiting light intensities we would need to determine the amount of photosynthetically usuable radiation (PUR). To do so we can integrate the product of PAR at each wavelength and the likelihood of absorption at that wavelength over the range of 400-750 nm. Since the action spectrum is plastic over time even in an individual organism, and certainly varies among organisms, we really can’t find one end-all be-all PUR value for a lamp that will be useful for different organisms, or even the same organism in different situations.
Having said that, typically the difference in intensity between lower Kelvn but brighter bulbs and higher Kelvin but dimmer bulbs is so great that the potential benefit of a more favorable spectrum is simply overwhelmed by the difference in intensity. But again, the difference in spectrum can only
potentially make a difference at low intensity. At higher intensity (near photosaturation) spectrum doesn’t matter at all. At that point there is so much light the efficiency of absorption is simply irrelevant.
Chris