Lecture III.1. Bacteria and Archaea. Prokaryotic and eukaryotic cells compared. Not every eukaryotic cell exhibits all features shown, e.g., animal and fungal cells lack plastids; fungal and most plant cells lack undulipodia, etc. 1 Prokaryotes. Small; mostly unicellular. No nucleus. DNA organized into 1. Circular chromosomes. 2. Plasmids – exchanged during conjugation or Plasmid transmission when taken up from envt. bacterial cells divide. Top. Inde- pendently. Bottom. As part of No mitosis; no meiosis. Re- the bacterial chromosome fol- produce by binary fission. lowing integration. No membrane-enclosed organelles – mitochondria, chloro- plasts, etc. But, bacteria do possess 1. Cytoskeleton proteins – some with eukaryotic counter- parts – so-called ESPs (Lecture II.4). 2. Protein-encased organelles such as carboxysomes that increase rate of photosynthesis. 2 Components of the Caulobacter Cytoskeleton. Caulobacter cells have homologs of each of the three major eukaryotic cytoskeletal systems. FtsZ localizes to the division plane and regulates cell division. MreB localizes to a dynamically contracting and expanding spiral and regulates cell shape, polarity & chromosome segregation. CreS (crescentin) localizes to Caulobacter’s inner curvature and regulates cell shape. 3 Carboxysomes concentrate CO , thereby facillitating its uptake by 2 RuBisCO. Left. Schematic representation of bacterial carbon fixation. Inorganic carbon is pumped into the cytosol as bicarbonate − ion (𝑯𝑪𝑶 ) and enters the carboxysome through small pores. The 𝟑 carboxysome contains carbonic anhydrase (CA), which converts bicarbonate into CO , and RuBisCO, which catalyzes the entry of 2 CO into the Calvin cycle, the latter via reaction with ribulose-1,5- 2 bisphosphate (RuBP). CO concentration within the carboxysome is 2 elevated by relative impermiability of the protein coat to CO . Right. 2 Schematic showing polyhedral structure of the protein coat. Colors correspond to different sheet paralogous proteins (products of duplicated genes) and interior enzymes RuBisCO and CA. Despite the similarity of the protein coat to the coats of viruses, there is no evidence as of this writing for phylogenetic relationships (Falkner et al. 2017. Nanoscale. 9: 10662-10673). From Samborska and Kimber (2012). 4 The entry of 𝐂𝐎 into the Calvin cycle is catalyzed by the enzyme 𝟐 RuBisCO. As the cycle turns, extra G3P molecules (bottom) are pro- duced. These three carbon sugars are subsequently converted into glucose (6 carbon sugar), which plants store as starch. 5 Two Prokaryote Domains: Archaea and Bacteria. Based on gene sequence of small ribosomal subunit. 1. Large and small subunits collaborate to assemble pro- teins (translation). 2. Highly conserved. 3. Three Domain theory re- quires that observations consistent with symbio- Translation entails passage of mRNA between the large and genic origin of Eukarya small ribosomal subunits, the result of lateral gene transfer. 6 Characters Shared by All Three Domains. Genetic 1. DNA the hereditary molecule. 2. Semi-conservative DNA replication – daughter mole- cules consist of one new strand; one old. 3. Transcription and translation: DNA RNA proteins. 4. “Universal” genetic code. Metabolic 1. Use of high-energy compounds such as ATP. 2. Production of ATP by glycolysis – anaerobic respiration. Structural 1. Ribosomes 2. Plasma membranes and flagella – but see below. 7 Table I. The Three Domains Compared. Character Bacteria Archaea Eukarya Archaea Similar to Bacteria Multicellular Rarely No Many Nuclear Mem- Rarely No Yes brane Circular Chromo- Most Yes No some Membrane–en- No No Yes closed organelles Sex No1 No1 Common Archaea Similar to Eukarya Histones associ- No Yes Yes ated with DNA Initiator tRNA FMet2 Methionine Methionine Archaea Unique Cell Membrane See accompanying figure Most include Cellulose Cell Wall No peptidoglycan peptidoglycan or chitin Flagella Grow from tip Grow from base Entirely different 1. DNA transfer and recombination. For Archaea, see Bernstein and Bern- stein. 2017. Pp. 103-117. In, Witzany (Ed.) Biocommunication of Archaea. Springer) 2. N-formylmethionine – methionine (sulfur containing amino acid) to which a (CHO) group has been added. 8 Archaea Present a Mix of Characters. Similar to Bacteria: 1. Unicellular. 2. No nuclear envelope 3. Circular chromosome 4. Absence of organelles Similar to Eukarya: 1. Histones associated with DNA. 2. Translation initiated by methionine (Met). Unique: 1. Plasma membrane structure. 2. Cell wall composition. 3. Flagellar structure and development – grow from base (archaea) vs. tip (bacteria). 9 Archaeal DNA is wrapped around histone tetramers. Shown here are electron micrographs taken of archaeal DNA in the presence of increasing concentrations of archaeal histone. From Marc et al. (2002). 10
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