The first plants

Who were the first plants?   At the risk of a scolding from some of my fellow botanists, I�m going to define plants broadly as organisms that use sunlight to make carbohydrates, and release oxygen gas as a byproduct -   in other words, organisms that photosynthesize.   There is in fact a diverse array of organisms that photosynthesize, including some bacteria and various kinds of algae that are only remotely related to one another; not to mention photosynthetic sea slugs!  So used in that way, "plant" is more of an ecological term for organisms that provide the base of the global food chain, not a precise taxonomic category. (see my post of October 5, 2011, "Plants and animals and kleptoplasts - oh my!")

My goal today is in fact to explore where photosynthesis came from, and we won't be distracted by taxonomic issues.  The first organisms to practice photosynthesis were bacteria, and the cyanobacteria were the first to employ modern photosynthesis, in which oxygen is released as a byproduct.  Bacteria are prokaryotic organisms that have a simple cell structure without nucleus or internal organelles.  They don�t have chloroplasts like true plants or algae, but their entire cell is adapted to conduct photosynthesis.  In my October 5, 2011  post, I described how cyanobacteria in fact were captured and domesticated to become the first chloroplasts.   I also argued as devil�s advocate that cyanobacteria are the only true plants � the multicellular organisms that we call plants are merely the luxury condominiums in which those captured cyanobacteria live!

Cyanobacteria come in many different forms and resemble true eukaryotic algae.  Mat-forming cyanobacteria
 that build stromatolites are similar to to A and B, in which cells are bound by a mucilaginous matrix. on the
surface of a rock.  Drawing from Haupt, Plant Morphology, McGraw-Hill, 1953.

Photosynthesis was invented only once, and passed on to various eukaryotic organisms as chloroplasts, which have been captured, stolen, and recaptured many times.  But when did it all start?  We travel to Australia, where we find some important clues.

In a few shallow, highly saline lagoons along the west coast of Australia, peculiar knobby pillars of rock called stromatolites stand like the disarrayed remnants of a terracotta army, eroded and distorted beyond recognition.  These monoliths were not carved by some ancient civilization, however, but are built up very slowly by microscopic living organisms.   On the tops of the knobs, you can find mats of living microorganisms, held together by a mucilaginous glue.  The glue is secreted primarily by cyanobacteria.  Many bacteria form mats like this.  It�s a good way to anchor yourself to a suitable location.  

Stromatolites at Shark Bay, Western Australia.  Photo by Paul
Harrison via Wikipedia.

Cyanobacteria are massively abundant organisms.  Many live as free-floating plankton or tangles of filaments, and account for as much as 50% of the photosynthesis occurring in open waters (Fig. 2). Others are attached to rocks as filaments or mats.   They are easily confused with true algae, which are eukaryotic organisms with nuclei, chloroplasts, and other organelles.  Before electron microscopy showed us the difference, cyanobacteria were called �blue-green algae.�

Microbial mats on the tops of stromatolites form deposits that extend the knobby pillars slowly upwards.  Particles of sediment and lime precipitated from the water get trapped in the sticky matrix, and periodically bury the living microbes.   The resourceful cyanobacteria in those instances migrate to the top of the sediment and begin a new mat.  This results in a fine structure of alternating light and dark bands.  Stromatolites are thus, like coral reefs, built by the living organisms that inhabit them.   There is a website devoted to Shark Bay in Western Australia (www.sharkbay.org) that includes facts, photos and a video swim through a grove of stromatolites.

             
Stromatolites turn out to be one of our most important clues as to the origin of plant life.  They have been around for about 3.5 million years.  Ancient fossilized stromatolites, which can also be found in parts of Australia, are in fact among the earliest signs of any kind of life on this planet.  They are abundant throughout much of the geological record, but became rather scarce around 500 million years ago.  This was the time of the  �Cambrian Explosion�, when many new kinds of animals appeared.  Stromatolites came under attack by voracious grazing animals equipped with hard, scraping mouth parts.  After that, they survived only in restricted sites too salty for such animals. 

Though there is still debate about whether the earliest stromatolites were formed by cyanobacteria or some earlier form of life, or even by some physical process, there is no doubt that they were building stromatolites  by 2.7 billion years ago � still a heck of a long ago!   Fossilized cells identifiable as cyanobacteria have also been found in slightly different types of rocks, the Warrawoona and Apex cherts of Australia, and these appear also to date back to 3.5 billion years ago.   Cyanobacteria were the plants, the photosynthetic organisms, that supported the Earth�s early ecosystem.  They did so virtually alone form almost 2 billion years, after which the first signs of eukaryotic algae began to appear.

The early photosynthesizers must have gradually built up enormous populations, for the oxygen they produced eventually transformed the vast oceans of our planet, and then the very atmosphere itself.  The scarcity of oxidized (�rusted�) minerals in the Earth�s oldest rocks (older than 3 billion years), indicates that there was very little free oxygen in the atmosphere at that time, so the oxidizing of crustal rocks is also evidence of plant life.  Iron is particularly abundant on Earth, and quite prone to rusting.  In the ancient seas there was a steady supply of iron bubbling up from underwater volcanic fissures, and from eroding surface rocks.  In its unoxidized state, iron is soluble in water, but when it oxidizes it forms insoluble molecules of hematite or magnetite, which sink to the bottom of the sea.  When oxygen became available in comparably huge quantities there were spectacular depositions of iron.  This resulted in distinctive and extensive rock layers known as the Banded Iron Formations.  The �rusting of the earth,� as it was called by Schopf (2006), is the source of most of the iron ore that is being ravenously consumed by modern civilization.

There is some evidence of limited iron formations about 3.5 billion years ago, but they did not become truly massive until the mid-2-billions.   This suggests that oxygen buildup may have occurred sporadically and slowly at first, but became overwhelming between 2.7 and 2.5 billion years ago.  Oxygen makes up 21% of our current atmosphere, but 20 times as much plant-generated oxygen may be tied up in banded iron formations.  The formation of iron deposits declined rapidly after about 2 billion years ago, as the supply of dissolved iron was depleted, and oxygen then began to build up in the atmosphere.   The transformed atmosphere made it possible for more complex organisms to evolve, leading to eukaryotic cells and the modern world of algae, plants, fungi and animals.

So cyanobacteria invented photosynthesis as we know it today, and were the first functional plants.  The story does not end there.  In the two billion years of their unchallenged domination of the Earth, cyanobacteria also invented the rudiments of aerobic respiration and nitrogen-fixation, two other essential metabolic processes - cyanobacteria had to do virtually everything themselves!  Aerobic respiration is the process by  which organisms �burn� carbohydrates to fuel their metabolic processes.  It is obviously essential  for animals, but also for plants.  How else could they  utilize the carbohydrate reserves that they produced for themselves?

In terms of starting ingredients, photosynthesis is easy.  All you  need are carbon dioxide and water, and both are abundant on the planet.  But to make certain other things, like protein, DNA, ATP and other vital organic molecules, you also need nitrogen, which isn�t so easy to come by.  Wait-a-minute, isn�t the Earth�s atmosphere about 70% nitrogen?  Yes, it is, but it�s nitrogen that is hard to use.  Nitrogen gas consists of two nitrogen atoms bound tightly together by a triple bond.  Most organisms can�t break those bonds to make use of the abundant nitrogen supply.  Enter the cyanobacteria.  Somewhere along the way they acquired the genes to split nitrogen, either through mutation and natural selection, or from some other bacterium that invented it first.   Bacteria freely share genes through the process of horizontal gene transfer, a process we exploit in genetic engineering.

Once the nitrogen molecule has been split, cyanobacteria are able to attach hydrogen atoms to the nitrogen, making ammonia, and ammonia then can become the amino group required to make the building blocks of protein � amino acids.  So cyanobacteria, arguably the most successful and certainly the most long-lived group of organisms, make carbohydrates, metabolize carbohydrates for energy, and make their own protein.  The rest of us have only stolen from them!

References:

Schopf, J. W. 2006.  Fossil evidence of Archaean life.  Phil. Trans. R. Soc. B 361: 869�885.

Website for images and more information about stromatolites in Australia: www.sharkbay.org  (under "Nature of Shark Bay")