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Global control: modulons • Different operons/regulons affected by same environmental signal – – – – – Presence of glucose Change from O2 to anaerobic growth Nitrogen limitation; phosphate starvation Oxidative stress Stationary phase; entering starvation state • Some methods of control: – alternate sigma factors; Sigma controls which promoters are used – cAMP and CRP 1 Bacterial response to environment • Rapid response crucial for survival – Simultaneous transcription and translation – Coordinate regulation in operons and regulons – Global genetic control through modulons • Bacteria respond to – – – – – Change from aerobic to anaerobic Presence/absence of glucose Amount of nutrients in general Presence of specific nutrients Population size 2 Quorum Sensing • Bacteria monitor their own population size – Pathogenesis: do not produce important molecules too soon to tip off the immune system. – Light production: a few bacteria make feeble glow, but ATP cost per cell remains high. – Bacteria form spores when in high numbers, avoid competition between each other. • System requirements – A signaling molecule that increases in concentration as the population increases; LMW – A receptor; activation of a set of genes 3 Chemotaxis and other taxes • Movement in response to environmental stimulus – – – – – Positive chemotaxis, attraction towards nutrients Negative: away from harmful chemicals Aerotaxis: motility in response to oxygen Phototaxis: motility to certain wavelengths of light Magnetotaxis: response to magnetic fields • Taxis is movement – Includes swimming through liquid using flagella – Swarming over surfaces with flagella – Gliding motility, requiring a surface to move over 4 Flagellar structures www.scu.edu/SCU/Departments/ BIOL/Flagella.jpg img.sparknotes.com/.../monera/ gifs/flagella.gif 5 Runs and Tumbles: bacteria find their way http://www.bgu.ac.il/~aflaloc/bioca/motil1.gif 6 Motility summarized • Flagella: protein appendages for swimming through liquid or across wet surfaces. • Axial filament: a bundle of internal flagella – Between cell membrane and outer membrane in spirochetes – Filament rotates, bacterium corkscrews through medium • Gliding – No visible structures, requires solid surface – Slime usually involved. 7 Axial filaments 8 http://images.google.com/imgres?imgurl=http://microvet.arizona.edu/Courses/MIC420/lecture_notes/spirochetes/gifs/spirochete_crossection.gif& imgrefurl=http://microvet.arizona.edu/Courses/MIC420/lecture_notes/spirochetes/spirochete_cr.html&h=302&w=400&sz=49&tbnid=BOVdHqe pF7UJ:&tbnh=90&tbnw=119&start=1&prev=/images%3Fq%3Daxial%2Bfilament%2Bbacteria%26hl%3Den%26lr%3D%26sa%3DG Gliding Motility Movement on a solid surface. Cells produce, move in slime trails. Cells glide in groups, singly, and can reverse directions. Unrelated organism glide: myxobacteria, flavobacteria, cyanobacteria; Recent data support polysaccharide synthesis, extrusion model. http://cmgm.stanford.edu/devbio/kaiserlab/about_myxo/about_myxococcus.html 9 Starvation Responses 10 • Bacteria frequently on verge of starvation – Rapid utilization of nutrients by community keeps nutrient supply low – Normal life typical of stationary phase – Bacteria monitor nutritional status and adjust through global genetic mechanisms • Types of responses – – – – Lower metabolic rates, smaller size (incr surface:volume) Induction of low Km uptake systems Release of extracellular enzymes, scavenging molecules Production of resting cells, spores Smaller size is better Increased surface to volume ratio Surf = 4 π r2 Vol = 4/3 π r3 Nutrients enter through cell surface; the more surface, the more nutrients can enter. Large interior means slow diffusion, long distances. The larger a sphere, the LOWER the surface/volume, creating “supply” problems to the cell’s interior. Smaller cell more easily maintained. 11 12 Uptake Rate Different Transport proteins Kinetics of Uptake systems low Km High Km system Low Km system high Km 0.000 0.100 0.200 0.300 0.400 0.500 0.600 [solute] 0.700 0.800 0.900 1.000 Bacteria switch to transport systems that work better at lower solute concentration. Extracellular molecules 13 • Enzymes – Polymers cannot enter cells – Proteins, starch, cellulose all valuable nutrients – Enzymes produced and released from the cell – LMW products taken up; nutrients gathered exceed energy costs. • Low molecular weight aids – Siderophores, hemolysins collect iron – Antibiotics may slow the growth of competition when nutrients are in short supply 14 Siderophores http://www.staff.uni-marburg.de/~oberthue/enterobactin.gif http://www-users.york.ac.uk/~srms500/research_group/pic_1.JPG Sporulation 15 • Resting cells – Cells respond to low nutrients by sporulation or slowing down metabolic rate, decr size. – Some cells change shape, develop thick coat – Endospores form within cells; very resistant. – Spores in bacteria generally are for survival • Not reproduction – A spore structure protects cells against drying, heat, etc. until better nutrient conditions return • An inactive cell can’t protect itself well Endospore formation Genetic cascade producing alternative sigma factors. http://www.microbe.org/art/endospore_cycle.jpg 16 Responses of microbes to hypertonicity 17 • If cell is in a hypertonic environment, water leaves the cell. Decrease of intracellular water causes proteins, etc. to precipitate out of solution, stop functioning. Bacteria respond by increasing the concentration of “compatible solutes” to partially balance the higher external solute concentration. http://www.unimarburg.de/fb17/fachgebiete/mikrobio/molmibi/forschung/ osmostress-response/image_preview 18 Compatible solutes • small neutral molecules accumulated in cytoplasm when external environment is hypertonic. • No net charge, not acidic or basic. http://www.thermera.com /images/Betaine.gif Stress proteins • Elevated temperatures turn on Heat shock proteins – Proteins help protect and repair other critical proteins in the cell – Heat and other environmental stresses turn on genes for these protective proteins. http://www.tulane.edu/~biochem/med/shock.gif 19