Bacteriorhodopsin phototrophy

Classification of the metabolic capabilities of microbes can be challenging. With few exceptions, macroorganisms are either photosynthesizing primary producers (photo-autotrophs) or consumers (organotrophs or more commonly, heterotrophs).

For microbes, the story is more complicated. In addition to phototrophy, microbes can be chemo- or litho-trophs meaning they are able to derive energy from the oxidation of inorganic compounds such as reduced sulfur. If they can use light or chemical energy to fix carbon, then they are considered autotrophs. If the energy they acquire can be used to synthesize ATP but not to fix carbon, they are dependent on external sources of organic carbon making them mixotrophs.

An example of a phototrophic mixotroph is pictured above. These are salt loving haloarchaea in salt production ponds near (in?) San Francisco. The red color is due to the transmembrane protein bacteriorhodopsin. Using this protein some haloarchaea can harness sunlight to pump protons across their cell membrane. This establishes a proton gradient across the membrane. This gradient can be used to generate ATP.

There is a large number of scientific papers on bacteriorhodopsin because of its relative simplicity, it has become a model system for the study of membrane associated ion pumps.

Image from here http://www.sfgate.com/cgi-bin/article.cgi?file=/chronicle/archive/2001/03/13/MN152047.DTL

These salt ponds are near San Francisco. If you want another view of the bay area ponds follow this link, select the satellite map and zoom way in. I tried this for a few of the other places I know these salt production ponds exist but the satellite images did not provide good enough resolution. An example is Bon Aire in the Netherlands Antillies

Cilliate kleptoplasty

Chloroplasts, like mitochondria, have their own chromosomal DNA. This is, of course, evidence for the endosymbiotic theory of the origin of chloroplasts. It is also useful because it allows researchers to use the DNA to identify the source of the chloroplasts present in kleptoplastic organisms. This is a fairly standard method and is the way that Gast et al 2007 determined the source of the chloroplasts in the Antartic dinoflagellates discussed in a previous post.

In a paper (pdf) a few years ago in Limnology and Oceanography, McManus et al. used the chloroplasts present in the kleptoplastic tide-pool ciliates, Strombidium oculatum and Strombidium styliferseen to help reveal an interesting life history. The chloroplasts were from the large multicellular macroalgae Enteromorpha clathrata which raised the question of how these unicellular cilliates were able to acquire macroalgal chloroplasts.

McManus et al. found that the cilliates don't appear to be grazing directly on the large strands of the mature algae but on the small motile reproductive cells, called zoospores, mature algal strands release.

In addition to chloroplasts, the zoospores contain a pigmented eyespot. As the photo above (from the paper's figure 1) demonstrates, the kleptoplastic cilliates contain a pigmented eyespot similar to the ones possessed by the zoospores. This suggests that the Strombidium cilliates also owe their phototaxic abilities to the alga cells they ingest.

Some other interesting points about these cilliates:

• They are tidal organisms and live by the rhythm of the tides, becoming active during low tide when tidal pools are calm, and then attach to surfaces and encyst during high tide, presumably to prevent them from being washed out to sea and away from their food.
  
• They appear to be obligate mixotrophs, unable to grow in the dark or in the absence of algal food.

Complete reference:
McManus, G. B., H. Zhang, and S. Lin. 2004. Marine planktonic ciliates that prey on macroalgae and enslave their chloroplasts. Limnol. Oceanogr. 49:308-313.

Kleptoplasty

Kleptoplasty is a wonderful term. It is used to describe the behavior of a group of organisms that are able to ingest algal cells and degrade the cells, but not the chloroplasts contained within the cells. The chloroplasts remain functional for some period of time during which the photosynthetic products generated by the sequestered chloroplasts are utilized by the new 'host'.

On the left is a figure from a paper by Gast et al. in the journal Enviornmental Microbiology from earlier this year showing a kleptoplastic dinoflagellate isolated from the Ross Sea in Antartica (the paper is in a free issue of the journal so go read the whole thing).

In addition to being a very interesting behavior from an ecological perspective, kleptoplasty is of evolutionary interest because the capacity to grow autotrophically by photosynthesis arose within dinoflagellates by the retention of chloroplasts from ingested algal cells. This ability appears to have arisen multiple times within dinoflagellates because not all contain chloroplasts from the same type of algal cell.

The name

I chose the name Mixotrophy because it suggests the blog will contain a varied diet of thoughts and ideas. Some original and some recycled.

Within biology, the term is used in at least two distinct ways. The more common usage (and the first one I encountered in my studies) describes a large, diverse group of photosynthetic organisms that are capable of acquiring energy from sunlight (autotrophy) and from the degradation of preformed organic compounds (heterotrophy). This definition from the Smithsonian Environmental Research Center gives an idea of how broadly the term is applied.

Mixotrophic organisms gain their nutrition through a combination of photosynthesis and uptake of dissolved or particulate organic material. However, they vary widely in their photosynthetic and heterotrophic capabilities. Some mixotrophs are mainly photosynthetic and only occasionally use an organic energy source. Others meet most of their nutritional demand by phagotrophy, but may use some of the products of photosynthesis from sequestered prey chloroplasts.

The other usage is limited to prokaryotes* and is much more specific in its meaning. Here, mixotrophy refers to organisms that are capable of acquiring energy from the oxidation of inorganic compound but are unable to fix carbon. This means they must obtain organic carbon for biosynthesis. A relatively well know example of an organism in this group is Beggiatoa


*Yes I am going to continue using the term and, at the risk of contributing nothing of substance to the discussion, I will probably put up a post about it later in the week