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Objective 3: TSWBAT recognize the processes by which bacteria respond to environmental changes by regulating transcription. © 2014 Pearson Education, Inc. Overview: Differential Expression of Genes Prokaryotes and eukaryotes alter gene expression in response to their changing environment Multicellular eukaryotes also develop and maintain multiple cell types Gene expression is often regulated at the transcription stage, but control at other stages is important, too © 2014 Pearson Education, Inc. Bacteria often respond to environmental change by regulating transcription Natural selection has favored bacteria that produce only the gene products needed by the cell A cell can regulate the production of enzymes by feedback inhibition or by gene regulation Gene expression in bacteria is controlled by a mechanism described as the operon model © 2014 Pearson Education, Inc. Figure 15.2 Precursor Feedback inhibition trpE gene Enzyme 1 trpD gene Enzyme 2 Regulation of gene expression trpC gene trpB gene Enzyme 3 trpA gene Tryptophan (a) Regulation of enzyme activity © 2014 Pearson Education, Inc. (b) Regulation of enzyme production Operons: The Basic Concept A group of functionally related genes can be coordinately controlled by a single “on-off switch” The regulatory “switch” is a segment of DNA called an operator usually positioned within the promoter An operon is the entire stretch of DNA that includes the operator, the promoter, and the genes that they control © 2014 Pearson Education, Inc. The operon can be switched off by a protein repressor The repressor prevents gene transcription by binding to the operator and blocking RNA polymerase The repressor is the product of a separate regulatory gene © 2014 Pearson Education, Inc. The repressor can be in an active or inactive form, depending on the presence of other molecules A corepressor is a molecule that cooperates with a repressor protein to switch an operon off For example, E. coli can synthesize the amino acid tryptophan © 2014 Pearson Education, Inc. By default the trp operon is on and the genes for tryptophan synthesis are transcribed When tryptophan is present, it binds to the trp repressor protein, which then turns the operon off The repressor is active only in the presence of its corepressor tryptophan; thus the trp operon is turned off (repressed) if tryptophan levels are high © 2014 Pearson Education, Inc. Figure 15.3 trp operon Promoter Promoter Genes of operon DNA trpE trpR Regulatory gene 3 mRNA Operator RNA Start codon polymerase mRNA 5 trpD trpC trpB trpA B A Stop codon 5 E Protein D C Inactive Polypeptide subunits that make up repressor enzymes for tryptophan synthesis (a) Tryptophan absent, repressor inactive, operon on DNA No RNA made mRNA Protein Active repressor Tryptophan (corepressor) (b) Tryptophan present, repressor active, operon off © 2014 Pearson Education, Inc. Figure 15.3a DNA trp operon Promoter Regulatory gene trpR mRNA Promoter Genes of operon trpE 3 trpD trpC trpB trpA B A Operator RNA Stop codon Start codon polymerase mRNA 5 5 E Protein Inactive repressor D Polypeptide subunits that make up enzymes for tryptophan synthesis (a) Tryptophan absent, repressor inactive, operon on © 2014 Pearson Education, Inc. C Figure 15.3b DNA No RNA made mRNA Protein Tryptophan (corepressor) Active repressor (b) Tryptophan present, repressor active, operon off © 2014 Pearson Education, Inc. Repressible and Inducible Operons: Two Types of Negative Gene Regulation A repressible operon is one that is usually on; binding of a repressor to the operator shuts off transcription The trp operon is a repressible operon An inducible operon is one that is usually off; a molecule called an inducer inactivates the repressor and turns on transcription © 2014 Pearson Education, Inc. The lac operon is an inducible operon and contains genes that code for enzymes used in the hydrolysis and metabolism of lactose By itself, the lac repressor is active and switches the lac operon off A molecule called an inducer inactivates the repressor to turn the lac operon on © 2014 Pearson Education, Inc. For the lac operon, the inducer is allolactose, formed from lactose that enters the cell Enzymes of the lactose pathway are called inducible enzymes Analogously, the enzymes for tryptophan synthesis are said to be repressible enzymes © 2014 Pearson Education, Inc. Video: lac Repressor Model Figure 15.4 Regulatory gene DNA Promoter Operator lacI IacZ No RNA made 3 RNA polymerase mRNA 5 Active repressor (a) Lactose absent, repressor active, operon off Protein lac operon DNA IacZ IacI IacY IacA Permease Transacetylase RNA polymerase 3 mRNA 5 Protein mRNA 5 -Galactosidase Inactive Allolactose repressor (inducer) (b) Lactose present, repressor inactive, operon on © 2014 Pearson Education, Inc. Figure 15.4a Regulatory gene DNA Operator lacI mRNA 5 Protein Promoter 3 RNA polymerase IacZ No RNA made Active repressor (a) Lactose absent, repressor active, operon off © 2014 Pearson Education, Inc. Figure 15.4b DNA lac operon IacI mRNA IacZ 3 5 Protein IacY IacA RNA polymerase mRNA 5 -Galactosidase Permease Transacetylase Inactive repressor Allolactose (inducer) (b) Lactose present, repressor inactive, operon on © 2014 Pearson Education, Inc. Inducible enzymes usually function in catabolic pathways; their synthesis is induced by a chemical signal Repressible enzymes usually function in anabolic pathways; their synthesis is repressed by high levels of the end product Regulation of the trp and lac operons involves negative control of genes because operons are switched off by the active form of the repressor © 2014 Pearson Education, Inc. Positive Gene Regulation E. coli will preferentially use glucose when it is present in the environment When glucose is scarce, CAP (catabolite activator protein) acts as an activator of transcription CAP is activated by binding with cyclic AMP (cAMP) Activated CAP attaches to the promoter of the lac operon and increases the affinity of RNA polymerase, thus accelerating transcription © 2014 Pearson Education, Inc. When glucose levels increase, CAP detaches from the lac operon, and transcription proceeds at a very low rate, even if lactose is present CAP helps regulate other operons that encode enzymes used in catabolic pathways © 2014 Pearson Education, Inc. Figure 15.5 Promoter DNA lacI IacZ CAP-binding site cAMP Operator RNA polymerase Active binds and transcribes CAP Inactive CAP Allolactose Inactive lac repressor (a) Lactose present, glucose scarce (cAMP level high): abundant lac mRNA synthesized Promoter DNA lacI CAP-binding site Inactive CAP IacZ Operator RNA polymerase less likely to bind Inactive lac repressor (b) Lactose present, glucose present (cAMP level low): little lac mRNA synthesized © 2014 Pearson Education, Inc. Figure 15.5a Promoter DNA lacI IacZ CAP-binding site cAMP Operator RNA polymerase Active binds and transcribes CAP Inactive CAP Allolactose Inactive lac repressor (a) Lactose present, glucose scarce (cAMP level high): abundant lac mRNA synthesized © 2014 Pearson Education, Inc. Figure 15.5b Promoter lacI DNA CAP-binding site Inactive CAP IacZ Operator RNA polymerase less likely to bind Inactive lac repressor (b) Lactose present, glucose present (cAMP level low): little lac mRNA synthesized © 2014 Pearson Education, Inc.