
This Is The Paper I Wrote on Endosymbiosis for Dr. Hettes' Cellular Biology Class, This really isn't that good because i wrote it in 3 hours :(
The evolution of our ancestry has long been a contested subject amongst biologists. Surely, the most interesting of topics has been the discussion on the endosymbiotic theory of organelles; a process that has incorporated various ubiquitous cellular ‘machinery’ into the cells of eukaryotes to perform highly specified jobs within the cell. However many theories are thrown about, the question remains: How did we end up with these small factories in our cells? Organelles such as the mitochondria and the chloroplasts are much too complex to have originated from the cell itself, or rather, originate autogenously, and many pieces of information point to the fact that it is highly unlikely that these organelles came from the cell itself. So, if not originating from the eukaryotic cells, where did these organelles come from? How did they manage to become incorporated into the cells for presumably a billion or so plus years? Other contested endosymbionts such as Peroxisomes, flagella, or even microtubules are perhaps too simple to have been from extracellular sources. What is the evidence behind these endosymbiotic organelles? These questions and many of the proposed theories are some of the most intriguing topics of discussion related to endosymbiotic theory.
While Lynn Margulis only introduced the proposed theory of endosymbiosis formally in recent decades, the idea was around long before Margulis’ time. The idea of a symbiotic relationship between two cellular organisms was first proposed by Konstantin Merezhkovski in 1905. His work in studying lichens (a composite organism that is symbiotic in nature), as well as contemporary ideas on cyanobacteria, sparked the idea of chloroplasts as a symbiotic component of plants cells (Margulis, 1993) . A man named Ivan Wallin proposed a similar idea in 1920, relating the idea that because mitochondria divide in cells much in the way bacteria do, and most of mitochondrial behavior is much like that of bacteria, postulating that mitochondria must be symbiotic within the eukaryotic cell. Margulis drew heavily on their work when writing her many books on the nature of endosymbiosis, and is recognized as the main proprietor of Endosymbiotic theory. Her integration of cellular biology, biochemistry, knowledge of the young Earth, and the theories behind life’s complex evolution have coalesced into a formal and unified theory of Eukaryotic origins, and gives us an idea of our own unicellular origins.
The most accepted forms of endosymbionts include mitochondria, and chloroplasts, which in structure and function most represent their bacterial ancestors. Mitochondria are found in virtually every eukaryotic cell, providing energy via synthesis of ATP from aerobic respiration, and are especially concentrated where needs for ATP is greatest. This process is very similar to prokaryotic bacteria, which derive energy from their environment by a very similar process. Chloroplasts are the organelles responsible for converting sunlight into energy via photosynthesis but this occurs in only some organisms such as plants, algae and bacteria (although not in Archaea) . The fact that this process occurs in photosynthetic bacteria, gave rise to the notion that chloroplasts had an extracellular origin, as first postulated by Merezhkovski. The most difficult organelles to comprehend as being endosymbionts are the peroxisomes and undulipodium. It is thought that peroxisomes are endosymbiotic with the cells in eukaryotes and are found in nearly every eukaryotic cell. The fact that peroxisomes are not part of the endomembrane system and the presence of similar proteins in the organelles suggests an endosymbiotic nature of the peroxisome. However, this notion may be disputed as synthesis of the peroxisome from the endoplasmic reticulum has been demonstrated and peroxisomes may in fact be part of the endomembrane system. Also, it should be noted that the similarity of proteins in many different peroxisomes across species should come as no surprise, as they all share a similar function. Undulipodium are the most interesting of the proposed endosymbionts. They are proposed to be from a group of bacteria called spirochetes. The spirochetes are a type of bacteria that have a complex of many microtubules that form flagella. However, in my own personal opinion, the integration of a completely separate organism is a whole lot more complicated than the simple two- protein ATP-ase that makes up flagellum. Flagella are little more than a rotor protein and a stator protein within an ATP-ase, with an attached microtubule to the rotor protein to serves as the driving force of the cell through its matrix.
The main theory of endosymbiosis is that at one time, proto-eukaryotic cells acquired proto-bacteria within their cellular matrix, which tended to be beneficial for both the proto-eukaryote and the proto-bacterium, as the proto-eukaryote gained a cellular machine that provided a specific function to the overall function of the cell, such as photosynthesis, or motility; and the proto-bacteria gained a living environment, and provided protection. Although, there are different mechanisms that are thought to have incorporated these proto-bacteria in the proto-eukaryotes. Serial endosymbiotic theory proposed the idea that these endosymbionts have been incorporated into the host cell one at a time, integrating these cells, and eventually the cells began to independently divide and continue on within the cell. The cells are thought to have phagocitzed other living cells for nutrients and that unsuccessful digestion of these cells could have led to a symbiotic relationship between the two. However, I find that there are many flaws with this proposal, even though it is the most supported theory of endosybiosis’s mechanisms. The fact that a cell that requires energy by phagocytizing other bacteria and it not being able to digest said bacteria, is one that is in poor shape and within the context of an early earth, these cells would probably not survive long. Also, there are no bacteria that actively use phagocytosis for predation (Margulis, 1993), and therefore the likelihood of these cells predating on these proto-bacteria is highly unlikely. All of these things together seem very unlikely, especially within the context of the evolving progression of life. It could be highly feasible that a few singularities of these events happening, but the entire basis of multi-cellular life depending on these rare events is improbable.
Another theory of endosymbiosis includes the autogenous theory of endosymbiosis. This idea postulates that the proto-eukaryotic cell itself formed its own organelles by invaginations of the plasma membrane and assorted proteins and enzymes came to work in these organelles developing specific jobs. This theory was hypothesized by David T. Smith and Cavalier in 1987, which was quickly pushed aside by Margulis’ theory a few years later. This theory is also unlikely because there is empirical evidence that mitochondria and chloroplasts have their own separate DNA, which highly resembles DNA of bacteria.
Yet there is another theory, somewhat newer than the rest, which plays with the idea that perhaps ancient mitochondria and chloroplasts were not actively engulfed by the host cell, but instead were parasitic bacteria that preyed upon the host cells, and do to mutual benefit of this symbiosis, stuck together through the ages.
The radical state of the early Earth had much to do with the formation and shaping of life on our planet, and therefore the constant flux of materials on earth had a hand in directing the evolutionary history of life. Most important (to us, anyway) is the beginnings of multicellular life. David Smith once noted “Many symbiotes are found in nutrient poor conditions or in hosts which depend on a nutrient poor diet” (Smith, 1987), which is probably true in respect to the formation of multicellular life.
During Earth’s early years, the atmosphere was full of noxious gases that most (if not all) multicellular life could not thrive in. However many ancient bacteria were very adept at converting noxious gases such as Carbon dioxide into oxygen by way of photosynthesis, these bacteria, like Cyanobacteria, thrived and converted vast amounts of carbon dioxide into diatomic oxygen. After many, many years of this process, the levels of oxygen rose sharply and levels of carbon dioxide decreased. Oxygen is actually a very corrosive element and is toxic to many organisms, especially ancient bacteria. Toxic levels of oxygen increased, and a carbon dioxide food source dwindled. There was hypothesized to be a mass die-off of organisms due to the toxic oxygen levels, but there were oxygen-loving organisms known as proto-mitochondria that thrived and survived in this new environment. Perhaps proto-mitochondria, were aerobic bacteria which oxidized fermentation products completely into carbon dioxide and water. But how did these protomitochondria end up in the larger cells? These proto-mitochondria are hypothesized to be parasitic ( this idea is backed up with the fact that many of the related bacteria to proto-mitochondria are in fact parasitic beings) and took advantage of the other weakened cells on early Earth. Cyanobacteria, the precursors to chloroplasts may have taken advantage of the waste products of the mitochondria to produce energy completely for the cell. The succession of mitochondria to chloroplasts also explains the fact that most all eukaryotic cells have mitochondria, whereas only those in the Plantae and some Fungi have the ability to photosynthesize. If an entire species of proto-mitochondria were parasitic, this would also help explain the large occurrence of these organelles, rather than singularities.
Sources
Ahmadijan, V.; Paracer, S. (1986), Symbiosis: an Introduction to Biological Associations, University Press of New England
Gray, M.; Doolittle, W.F. (1982), Has the Endosymbiont Hypothesis Been Proven?, Dalhousie University, MICROBIOLOGICAL REVIEWS, Retrieved from:
http://mmbr.asm.org/cgi/reprint/46/1/1?view=long&pmid=6178009
On March 23, 2009
Lane, N. (2006), Power, Sex, Suicide: Mitochondria and the Meaning of Life, Oxford University Press.
Margulis, L. (1993), Symbiosis in Cell Evolution, W.H. Freemen Company Press
Oxford University (2008), Oxford Dictionary of Biology, Oxford University Press.
Smith, D.C., Douglas, A.E. (1987), The Biology of Symbiosis, Edward Arnold (Publishers) Ltd.
Thomas, L. (1974), The Lives Of a Cell, The Viking Press Inc.
Tzagoloff, A. (1982), Mitochondria, Plenum Press