Endosymbiont - Endosymbiosis Theory and Mitochondria and Chloroplasts

Endosymbiosis Theory and Mitochondria and Chloroplasts

The endosymbiosis theory attempts to explain the origins of organelles such as mitochondria and chloroplasts in eukaryotic cells. The theory proposes that chloroplasts and mitochondria evolved from certain types of bacteria that eukaryotic cells engulfed through endophagocytosis. These cells and the bacteria trapped inside them entered a symbiotic relationship, a close association between different types of organisms over an extended time. However, more specifically, the relationship was endosymbiotic, meaning that one of the organisms (the bacteria) lived within the other (the eukaryotic cells).

According to endosymbiosis theory, an anaerobic cell probably ingested an aerobic bacterium but failed to digest it. The aerobic bacterium flourished within the cell because the cell's cytoplasm was abundant in half-digested food molecules. The bacterium digested these molecules with oxygen and gained great amounts of energy. Because the bacterium had so much energy, it probably leaked some of it as Adenosine triphosphate into the cell's cytoplasm. This benefited the anaerobic cell because it enabled it to digest food aerobically. Eventually, the aerobic bacterium could no longer live independently from the cell, and it therefore became a mitochondrion. The origin of the chloroplast is very similar to that of the mitochondrion. A cell must have captured a photosynthetic cyanobacterium and failed to digest it. The cyanobacterium thrived in the cell and eventually evolved into the first chloroplast. Other eukaryotic organelles may have also evolved through endosymbiosis; it has been proposed that cilia, flagella, centrioles, and microtubules may have originated from a symbiosis between a Spirochaete bacterium and an early eukaryotic cell, but this is not widely accepted among biologists.

There are several examples of evidence that support endosymbiosis theory. Mitochondria and chloroplasts contain their own small supply of DNA, which may be remnants of the genome the organelles had when they were independent aerobic bacteria. The single most convincing evidence of the descent of organelles from bacteria is the position of mitochondria and plastid DNA sequences in phylogenetic trees of bacteria. Mitochondria have sequences that clearly indicate origin from a group of bacteria called the alphaproteobacteria. Plastids have DNA sequences that indicate origin from the cyanobacteria (blue-green algae). In addition, there are organisms alive today, called living intermediates, that are in a similar endosymbiotic condition to the prokaryotic cells and the aerobic bacteria. Living intermediates show that the evolution proposed by the endosymbiont theory is possible. For example, the giant amoeba Pelomyxa lacks mitochondria but has aerobic bacteria that carry out a similar role. A variety of corals, clams, snails, and one species of Paramecium permanently host algae in their cells. Many of the insect endosymbionts have been shown to have ancient associations with their hosts, involving strictly vertical inheritance. In addition, these insect symbionts have similar patterns of genome evolution to those found in true organelles: genome reduction, rapid rates of gene evolution, and bias in nucleotide base composition favoring adenine and thymine, at the expense of guanine and cytosine.

Further evidence of endosymbiosis are the prokaryotic ribosomes found within chloroplasts and mitochondria as well as the double-membrane enclosing them. It used to be widely assumed that the inner membrane is the original membrane of the once independent prokaryote, while the outer one is the food vacuole (phagosomal membrane) it was enclosed in initially. However, this view neglects the fact that i) both modern cyanobacteria and alpha-proteobacteria are Gram negative bacteria, which are surrounded by double membranes; ii) the outer membranes of the endosymbiotic organelles (chloroplasts and mitochondria) are very similar to those of these bacteria in their lipid and protein compositions. Accumulating biochemical data strongly suggest that the double membrane enclosing chloroplasts and mitochondria derived from those of the ancestral bacteria, and the phagosomal membrane disappeared during organelle evolution. Triple or quadruple membranes are found among certain algae, probably resulting from repeated endosymbiosis (although little else was retained of the engulfed cell).

These modern organisms with endosymbiotic relationships with aerobic bacteria have verified the endosymbiotic theory, which explains the origin of mitochondria and chloroplasts from bacteria. Researchers in molecular and evolutionary biology no longer question this theory, although some of the details, such as the mechanisms for loss of genes from organelles to host nuclear genomes, are still being worked out.