Fiddling with photosynthesis (Alien Plants IV)

'Alien Plants IV'? Where are the other 'alien plant' posts? Well, 'Alien plants I' and 'II' were published a long time ago, and 'Alien Plants III' was not labelled as such: that would be the post 'The black, black grass of home...' posted one year ago. That one was more serious than the first two, and  dealt with the colour of plants on Earth. To be succinct: green does not equal photosynthesis.

Click to enlarge. Copyright 2007 University of Chicago. From: Nature's palette by David Lee

As can be seen from the absorption spectrum of chlorophyll above, photosynthesis does not use the green portion of the spectrum, so that portion gets reflected for us to see. In doing so plants ignore much energy potentially available to them, as green is right in the part of the spectrum where the sun emits a lot of light. You might think that photosynthesis would evolve to make the most of the light falling on it, and, if so, you would predict that Earth plants should be purple (see the 'black grass'  post for speculations why some bacteria are purple but plants are not).

Some people wonder whether we can predict the colour of plants on a planet by looking at the spectrum of its sun. Earth's example definitely suggests that we cannot, so I personally see no problems with filling hypothetical planets with plants of just about any colour; well, as long as the absorbed colour is present in that sun's spectrum, of course. A perfect photosynthesis process would be able to use light of every frequency equally well, with the effect that such plants would be grey or black.

After writing the 'black grass' post I returned to the question why it is difficult to come up with alien-looking plants. Intuition suggested that there would be only so much you could do with plant shapes: flat leaves fixed to the ends of a branching structure seem so sensible that they are probably universal, so plants everywhere would look similar. Perhaps so, but intuition is not a reliable predictor in science, so some old-fashioned studying was called for. I recommend 'The Life of a Leaf' by Steven Vogel, who also wrote a fine book on biomechanics.

The fun part will be designing new plant shapes, if possible, but before we get to that there is some work to do, I am afraid. This post starts with photosynthesis on Earth, to find out if it can be tweaked to produce plants with a high degree of 'alienosity'.

Click to enlarge; Based on Long SP et al. Can improvement in photosynthesis increase crop yields? Plant, cell , and environment 2006; 29: 315-330

1. Efficiency of photosynthesis
The job of photosynthesis is to take water, CO2 and light, and turn out carbohydrates to use as energy sources and building materials, with O2 as a leftover waste product. Although the total energy capture by photosynthesis outranks human power consumption by far, photosynthesis is less efficient than the photovoltaic process used in solar panels. Photosynthesis is surprisingly inefficient.  The image above is based on analyses done by scientists looking for ways to improve crop yield. The 'black grass' post explained that only a portion of sunlight is used for photosynthesis, and the papers show that portion to be about half of the available energy. The graph above states the efficiency of each step, which is which fraction of energy gets passed on to the next step. The efficiency of the first step is 0.5: of 100% light to start with, 50% is left. That's a big loss. 
   The efficiency of the next step is 0.9. In terms of the original amount of light 45% goes on to the next step. And so it goes on, multiplying all the efficiency factors in turn, step by step, until only about 5% of the original energy is left at the end. As I said, not impressive at all. I should add that this holds for the so-called C3 photosynthesis type. The C4 type does better, managing to end up at 6 to 6.5%.  That does not seem like a big improvement, but it is still up to 30% better than C3 photosynthesis.      

One biochemical step deserves additional mention: 'photorespiration'. The reactions that take in H2O, CO2 and light to turn them into sugars and O2 are not exactly simple; an important enzyme capturing CO2 is ribulose-1,5-bisphosphate carboxylase oxygenase (no wonder that it is called 'Rubisco'). Rubisco deserves to be known, if only because it is probably the most common protein on Earth. Its job is to speed up the reaction binding CO2 that ultimately ends in O2. Oddly, Rubisco binds quite readily with O2, driving a process in the wrong direction! This backwards process is called 'photorespiration' and has puzzled biologists a lot. Its presence suggested that it might have some use, but apparently plants do quite well in artificial atmospheres without any O2 at all, so photorespiration seems to be a gigantic and puzzling waste.

2. Bright light: photosynthesis saturation
As if the above series of limitations is not enough, there is another one: photosynthesis saturates. Photosynthesis normally increases with the level of light but only up to a point. If light intensity increases beyond that point, photosynthesis cannot increase with it (it may apparently even decrease to protect the plant). Whether this is an important limitation depends on where you are: to catching the maximum amount of light to reach the Earth's surface, you will have to stand at the equator, at noon, on a clear day. The C3 type of photosynthesis can only use about a quarter of the light there! If you were to add that step to the image above, the scheme would start with a giant loss of 75% right at the start. Seen in that light (pun intended) the overall efficiency of 5% becomes an even less impressive 1.25%.

Then again, it is a bit unfair to set light at noon in the tropics on a cloudless day as the standard. Living at higher latitudes, clouds and shadows from mountains or leaves will limit the amount of light that reaches a plant, so in many cases the saturation point will never be reached. That is fine for those plants, but the tropics are still there, and photosynthesis could do a lot more for tropical plants  if their saturation point would lie at a higher intensity.

3. Shadows: the photosynthesis compensation point      
Plant cells burn molecules with the help of oxygen to free stored energy and use that for their metabolic needs, exactly like animal cells. This process is called cellular respiration and does the opposite of photosynthesis. As the amount of light decreases, photosynthesis will be less effective and produce less oxygen, while cellular respiration keeps using it a stable rate. At some shadowy light intensity the two processes are matched: the compensation point. When light levels drop beyond that point, plants become net users of oxygen and energy instead of producers. Plants can survive that state and in fact do so every night, but over time there must be a net profit. There are many places, such as the floor of dense forests, where it permanently too dark for photosynthesis to work.

"It's photosynthesis, Jim, but not photosynthesis as we know it". 
With all this in mind there seems to be ample opportunity to tinker with the process and design an alien photosynthesis. Mind you, photosynthesis could well be even less efficient on an alien planet than on Earth, and that possibility should not be dismissed out of hand. World builders have a strong tendency to design super-organisms, better than what Earth has to offer, but that is not very realistic. For once I will follow the flow and aim to improve on Earth's state of affairs. The following list concerns my suggestions how to improve on off-the-shelf photosynthesis:

Alien photosynthetic to-do list
- Have your photosynthesis process use a larger portion of the light falling on it
- Increase its affinity for CO2 (abolish photorespiration!) and improve reaction speed 
- Increase its saturation point so it can use intense light
- Lower the compensation point so it can work with less light.    

This 'to-do list' assumes that there are numerous biochemical pathways that can take in CO2, H2O and light and produce carbohydrates. Such processes may be centred on completely different pigments, sensitive to other wavelengths.

Click to enlarge; copyright University of Chicago. From: Nature's palette by David Lee

The illustration above has nothing to do with photosynthesis itself, but illustrates that there are many pigments in vision that are sensitive to varying wavelengths and to varying ranges of wavelengths. The pigment of the nectar-varying bat is interesting in that it is sensitive to a very broad range of light with a broad peak in the green area. A pigment like that, used for photosynthesis, would result in plants using light best where there is most of it, without throwing the rest away. Such plants would probably be a boring dark purplish grey.   

You may well ask whether all this biochemical tinkering will make plants look different. If they still look like Earth plants but grow faster the exercise loses much of its appeal, doesn't it? I think they would look different: if leaves can use all light falling on them, that will have consequences for any leaves underneath; simple blobs or needles might replace complex leaves; the ability to have fewer leaves might induce trees to grow higher; plants might continue to grow through winter, etc., etc.

Click to enlarge; by Ghedoghedo

Of course, apart from biochemistry different biomechanical design principles will also result in differently looking plants. To see whether that approach yields interesting choices, we may need to travel back to the Silurian and Devonian and have a look at designs principles that came into being when land plants first struggled against gravity. Changing designs and changing plant biochemistry ought to result in enough 'alienosity' to please anyone. We'll see...

The black, black grass of home...

Black? I hoped that by substituting 'green' with 'black' in the title of this evergreen ('everblack'?), your mind would create an image in which plants are suddenly no longer green but black. Plants hardly ever feature as more than background material in science fiction. SF artists may try to come up with odd plant shapes, but the general colour of your generic SF plant is green. Personally, I also often used green plants in my Furahan paintings without conscious thought.

Click to enlarge; copyright Gert van Dijk

What you see here are 'blackgrasses' on Furaha. In effect, they are largely brownish, but in this case at least I did not fall for the 'plants are green' trap. There are exceptions to ubiquitous greenery though, and the best-known one is probably Well's Martian 'red weed'. Anyway, perhaps it is time to think a bit harder about the colour of plants, first on Earth, then elsewhere. This post will be a bit technical; sorry for that. For more thoughts on this issue, see here and here.

The 'green, green grass of Earth' may trigger associations chlorophyll and photosynthesis ('chloro' means 'green' and 'phyll' is derived from 'leaf' in classical Greek, so it means 'green leaf stuff'). Photosynthesis concerns the trick of capturing the energy in light and transferring it to chemical energy (ATP), and chlorophyll is at the centre of that trick: it captures a photon, setting loose an electron that sets a cascade of motions going. As Chlorophyll is green, you might think that a green colour is good for photosynthesis. It is not: green light is almost useless for photosynthesis using chlorophyll.

Click to enlarge; from Wikipedia

Remember that what we call white light is a composite of a range of wavelengths in the electromagnetic spectrum, ranging from deep purple through blue, green, yellow and red to deep red. It is no coincidence that we call that portion 'visible light'. When light falls on an object some wavelengths are absorbed, while others are reflected. If an object looks green to us, that means that green light is reflected, meaning it is NOT used by photosynthesis. Above is an image of the absorption spectrum of two chlorophyll variants. A peak at a specific wavelength means that light at that wavelength is absorbed and used by chlorophyll. There are peaks in the red and blue parts of the spectrum, but not in the green portion of the spectrum. Does that matter? The answer depends on whether there is in fact a lot of light in that part of the spectrum, so let's compare the absorption spectrum of chlorophyll with the light output of the sun.

Click to enlarge; from Wikipedia

The atmosphere selectively absorbs some wavelengths, so what is relevant is how much of each wavelength reaches the Earth's surface. That is shown above. Try to find the visible part of the spectrum , from about 350 to 750 nanometer. You will see that there is a lot of light there.

Click to enlarge; from here

Here is a graph combining all the previous information. Note that the title states that chlorphyl is well-adapted to use solar energy. Well, yes, in the sense that it is roughly sensitive to light in the area where there is most energy. However, there is this big conspicuous gap, meaning that lots of light is unused by plants, mostly of the green variety. In fact, Earth plants would get on quite well if the sun did not emit all that energy at green wavelengths. (We would not like that though, as there would be about 40% less light to see with, and our ability to see details would be harmed as that depends to a large extent on green light; but that is another matter).

All this suggests that chlorophyll is not the best of all possible light absorbers on Earth. To make the most of sunlight, you would want a molecule that is responsive to a much broader part of the spectrum. Such a molecule would reflect very little light, so it would be black or at least very dark. You can also argue that, if chlorophyll can get away with using just part of the spectrum, so could another molecule. In fact, chlorophyll is not the only molecule used in photosynthesis.

Click to enlarge; taken from this site

'Bacteriorhodopsin' is a well-known example, occurring in some bacteria. Above is a graph in which its spectrum is overlaid on that of chlorophyll. This pigment has a rather broad absorption spectrum, but with a peak at precisely the spot where you would want it, meaning where the sun puts out much light: in the green / yellow parts of the spectrum. As a result, it looks reddish. There is a theory that these bacteria formed mats overlying the very first plants. The only light they let through was at wavelengths the bacteria did not use, so chlorophyll evolved to pick up just those wavelengths. This is a fascinating idea, but one thing that has me worried is the following: if chlorophyll could evolve such a detailed sensitivity to particular wavelengths, why did it stop evolving once plants outgrew those bacteria? Wouldn't it have made sense to retune it afterwards? If it would be sensitive to green light as well we would have black plants, and if it would be mostly responsive to green light plants would be purple.

I have no idea how easily evolution can tinker with light-sensitive molecules to shift their absorption spectrum. Still, light-sensitive pigments occur in vision as well as in photosynthesis, and there is an astonishing variety of pigments for colour vision in the animal kingdom. It seems that such pigments can evolve readily. But chlorophyll seems just to be sitting there, blind to all that glorious green light. Adding insult to injury, the chemical steps following chlorophyll are also singularly inefficient. Part of the problem seems to be that there is very little CO2 in the atmosphere, and that the molecule that takes in CO2 to strip the carbon from it leaving O2, quite readily works in the wrong direction, so the work is partly undone. After all this, the calculated efficiency of chlorophyll photosynthesis is on the order of 4-7%. Seeing how life on Earth depends on photosynthesis, that is a bit worrying.

In summary, the blindness of chlorophyll to green light suggests that other molecules with a different absorption spectrum could be just as efficient (or inefficient). It also suggests that you cannot tell the colour of plants well from the spectrum of the sun in their solar system. If you apply that reasoning on Earth, you will proclaim that Earth's plants are purple...

click to enlarge; copyright Chris Webb or Scientific American

In order to show something besides graphs, I will show a nice image used on the cover of Scientific American. I found it on the website of the artist (Chris Webb) right here.

In another post I may go into the consequences of all this for alien plants. For now, just consider some parallel evolution going on, resulting in different groups of 'plants' using different pigments. Forests on such planets need not show shades of green only, but could sport a riot of colours. Ah! The green, blue, yellow, purple grass of home...