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For a typical small molecule e.
Bacteria that do induce currents for their benefit e. Another postulated function of primitive motility, swimming for the sake of running into more molecules, also does not work: Thus, if diffusion of molecules into the cell is the only matter of concern, a bacterium will do just as well by sitting still as it will by stirring or swimming.
The reason bacteria swim is not to increase diffusion but to find locations with a higher local concentration of nutrient molecules Purcell, ; Berg, ; Vogel, However, a typical value for a is 0. Thus, fundamental physical considerations make the hypothesized stirring filament an unlikely intermediate.
Similarly, Rizzotti implies that the F0-c subunit is homologous with the flagellar motor proteins MotAB, but sequence homology has instead been discovered homology between MotAB and a phylogenetically widespread family of proteins that couple protonmotive force to diverse membrane transport processes.
The homologies could be explained by invoking additional independent cooption events, but this would require a rather more complex scenario than that presented by Rizzotti. Therefore several of the ideas proposed here have been previously raised in informal debates about flagellar evolution.
Millerand Musgrave review this aspect of the debate in detail, and Musgrave proposes a model that is similar in outline to that presented here, although his account is more general. Phylogenetic context and assumed starting organism The paradigm for prokaryote phylogeny, if there is one, is the universal rRNA tree.
This shows a number of widely separated bacterial lineages, with archaea and eukaryotes separated from them all by a very long branch. This tree is unrooted, and many possible rootings have been proposed in the literature.
As these are the most remote and difficult phylogenetic events it is possible to study, and as there is by definition no outgroup to life in general, the debate can be expected to continue for some time. For current purposes the most important point is that flagella are widespread across the bacterial phylogenetic tree, with losses in various taxa and no clearly primitive nonflagellate taxa.
It is therefore assumed that flagella evolved near the base of the bacterial tree. Rizzotti and others e. However, the very general consideration that most of the bacterial phyla are gram negative, including the many different taxa that come out as basal on different analyses, weighs against this hypothesis.
Therefore, we shall side with Cavalier-Smith, who argues that the last common ancestor was gram-negative. He has put forward the most detailed model for the origin of bacteria and the double membrane Cavalier-Smith, a, a.
The model thus begins with a generic double-membraned, gram-negative bacterium. The present model will begin with a reasonably complex bacterium, already possessing the general secretory pathway and type II secretion system, as well as signal transduction, a peptidoglycan cell wall, and F1F0-ATP synthetase.
Cavalier-Smith a hypothesizes that chlorobacteria may be the most basal offshoot of the tree and be primitively nonflagellate. An export system plus a mechanism to cross the outer membrane forms a secretion system.
Bacteria make use of a number of distinct secretion systems, reviewed as a group elsewhere Hueck, ; Thanassi and Hultgren, a; van Wely et al. Six major well-characterized secretion systems Figure 4aFigure 5 are reviewed by Thanassi and Hultgren a.
It is likely that systems will be added to the list in time. Systems with components homologous to flagellar components.
For components with well-documented homology to flagellar components, the name according to the unified nomenclature for type III secretion systems proposed by Hueck is given Sct: Secretion and Cellular Translocation first, followed by the currently accepted name for the Hrp protein.
The name of the flagellar homolog is shown in brackets. Further possible homologies are discussed in the text. Various secretion systems of prokaryotes. Based on several sources Jarrell et al. Another nucleotide may be substituted for ATP in some cases. See Table 4 for description of the functions of the systems.
About 10 well-conserved protein species make up the core of the type III export apparatus, which is used to export the axial components of bacterial flagella rod, hook, filament, adaptor, and cap proteins. In it was discovered that homologs of these proteins are also used to secret virulence factors in a diverse array of proteobacterial pathogens, such as Yersinia pestis, Salmonella typhimurium, Pseudomonas aeruginosa and enteropathogenic E.
This includes the two currently known members virulence and flagellar secretion systems and any unknown homologs. The existence of a nonflagellar type III export apparatus falsifies the argument that flagellar components are useless if they are not part of a fully functioning flagellum.
Thus, the model for the origin of flagella begins with the hypothesis of a primitive type III export system. This hypothesis, however, requires justification on several grounds in order to ameliorate obvious objections.eilean ni chuilleanain essay about myself paper ptsd research bjm tu essay essayalliance essays about beta club emblem middlebury supplement essay role of individual.
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