Ch. 18 Regula'on of Gene Expression BIOL 222
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1 Ch. 18 Regula'on of Gene Expression BIOL 222 Overview: Conduc'ng the Gene'c Orchestra Prokaryotes and eukaryotes alter gene expression in response to their changing environment In eukaryotes gene expression dictates cell molecules play many roles in gene expression in eukaryotes Bacterial Transcrip'onal Regula'on Natural has favored bacteria that produce only the products needed by that cell A cell can regulate the produc@on of enzymes by feedback inhibi@on or by Feedback inhibition Precursor Enzyme 1 Enzyme 2 Enzyme 3 trpe gene trpd gene trpc gene trpb gene trpa gene Regulation of gene expression gene regula@on Gene expression in bacteria Tryptophan (a) Regulation of enzyme activity (b) Regulation of enzyme production controlled by the operon model 1
2 Operons: The Basic Concept A cluster of func@onally related genes can be under coordinated control by a single on- off switch operator The regulatory switch operon usually posi@oned within the promoter en@re stretch of that includes operator, the promoter, and the genes that they control Operons: The Basic Concept that switches operon off prevents gene transcrip@on binds to operator, blocking polymerase product of a separate regulatory gene co can be ac@ve or inac@ve depending on presence of other molecules cooperates with a protein to switch an operon off For example, E. coli can synthesize the amino acid tryptophan Operon Repression By default the trp operon is on genes for tryptophan synthesis are transcribed When tryptophan is present it binds to the trp protein Together turning the operon off By both binding to regulator is ac@ve only in the presence of its co tryptophan thus the trp operon is turned off (repressed) if tryptophan levels are high 2
3 Fig Inactive trp operon (a) Tryptophan absent, inactive, operon on Genes of operon trpr trpe trpd trpc trpb trpa Regulatory Operator gene 3ʹ Start codon Stop codon m m 5ʹ 5ʹ polymerase E D C B A Polypeptide subunits that make up enzymes for tryptophan synthesis No made m Tryptophan (co) Active (b) Tryptophan present, active, operon off Two Types of Nega've Gene Regula'on Repressible operon usually on binding of a to the operator shuts off transcrip@on Like trp operon Inducible operon usually off inducer inac@vates the turns on transcrip@on Inducible Operon lac operon inducible operon Genes code for enzymes For hydrolysis and metabolism of lactose By itself, the lac is ac@ve switches the lac operon off inducer inac@vates the to turn the lac operon on 3
4 Fig Regulatory gene Operator laci lacz No 3ʹ made m 5ʹ polymerase PLAY Active (a) Lactose absent, active, operon off lac operon laci lacz lacy laca m 5ʹ 3ʹ polymerase m 5ʹ β-galactosidase Permease Transacetylase Allolactose (inducer) Inactive (b) Lactose present, inactive, operon on Operon Inducible enzymes usually func@on in catabolic pathways synthesis is induced by a chemical signal Repressible enzymes usually func@on in anabolic pathways synthesis is repressed by high levels of the end product Regula@on of the trp and lac operons involves nega%ve control of genes because operons are switched off by the ac.ve form of the Posi've Gene Regula'on Ac'vator (accelerator) Some operons subject to posi@ve control through a s@mulatory protein Such as catabolite ac@vator protein (CAP) When glucose (a preferred food source of E. coli) is scarce CAP is ac@vated by binding with cyclic AMP Ac@vated CAP binds promoter of lac operon increases affinity of polymerase accelera@ng transcrip@on When glucose levels increase CAP detaches from the lac operon transcrip@on returns to a normal rate 4
5 Fig laci CAP-binding site camp Active CAP Operator lacz polymerase binds and transcribes laci CAP-binding site Inactive CAP Inactive CAP Allolactose Inactive lac (a) Lactose present, glucose scarce (camp level high): abundant lac m synthesized Operator lacz polymerase less likely to bind Inactive lac (b) Lactose present, glucose present (camp level low): little lac m synthesized Differen'al Gene Expression in Eukaryotes Signal Cells in a mul@cellular eukaryo@c organism gene@cally iden@cal differen'al gene expression Creates differences between cell types Cap NUCLEUS Chromatin Chromatin modification Gene available for transcription Gene Transcription Primary transcript Intron processing Tail m in nucleus Transport to cytoplasm CYTOPLASM m in cytoplasm Degradation of m Translation Polypeptide processing Degradation of protein Active protein Transport to cellular destination Cellular function Regula'on of Chroma'n Structure highly packed heterochroma@n Genes are usually not expressed Chemical modifica@ons to histones and of chroma@n influence both chroma@n structure and gene expression 5
6 histone acetyla'on Histone Modifica'ons acetyl groups are axached to charged lysines in histone tails loosens structure histone methyla'on can condense Can cause long term gene Leading to altered cell double helix (a) Histone tails protrude outward from a nucleosome Histone tails Amino acids available for chemical modification phosphoryla'on next to a methylated amino acid can loosen chroma@n Unacetylated histones Acetylated histones (b) Acetylation of histone tails promotes loose chromatin structure that permits transcription Epigene'c Inheritance epigene'c inheritance traits transmixed by mechanisms not directly involving the nucleo@de sequence Due to methyla@on, acetyla@on, or phosphoryla@on Organiza'on of a Typical Eukaryo'c Gene control elements segments of noncoding help regulate transcrip@on by binding certain proteins cri@cal to the precise regula@on of gene expression in different cell types Enhancer (distal control elements) Proximal control elements Poly-A signal sequence Termination region Upstream Intron Intron Transcription Downstream Primary transcript 5ʹ Intron Intron processing Cleaved 3ʹ end of primary transcript Intron Poly-A signal Coding segment m 5ʹ Cap 5ʹ UTR Start codon Stop codon 3ʹ UTR Poly-A tail 3ʹ 6
7 Enhancers and Specific Transcrip'on Factors Proximal control elements located close to the promoter Distal control elements Activators Distal control Enhancer element TATA box Gene enhancers PLAY -bending protein General transcription factors Group of mediator proteins binds to an enhancer polymerase II of a polymerase II gene Transcription initiation complex synthesis Processing Post- transla@onal control alterna've splicing different m molecules from the same primary transcript depending on which exons are used Primary transcript m s Troponin T gene splicing or m Degrada'on Life span of m molecules key to determining protein synthesis determined in part by sequences in the leader and trailer regions Eukaryo@c m more long lived than prokaryo@c m Methylated cap, Poly- A tail 7
8 Processing and Degrada'on protein processing The cleavage and/or of chemical groups subject to control Proteasomes giant protein complexes bind protein molecules and degrade them Ubiquitin Proteasome Proteasome and ubiquitin to be recycled to be degraded Ubiquitinated protein entering a proteasome fragments (peptides) Ini'a'on of Transla'on Other forms of regula@on include regula@on of the ini@a@on of transla@on of selected ms can be blocked by regulatory proteins in the cytosol that bind to sequences or structures of the m Alterna@vely, transla@on of all ms in a cell may be regulated simultaneously Ie. transla@on ini@a@on factors are simultaneously ac@vated in an egg following fer@liza@on Noncoding s Only a small frac@on of codes for proteins, r, and t A significant amount of the genome may be transcribed into noncoding s Noncoding s regulate gene expression at two points chroma@n configura@on m transla@on 8
9 Micros and Small Interfering s Micros (mis) small single- stranded molecules Hairpin mi Hydrogen bond bind to m Dicer can degrade m or block its transla@on 5ʹ 3ʹ (a) Primary mi transcript mi miprotein complex m degraded Translation blocked (b) Generation and function of mis Micros and Small Interfering s interference (i) Process of inhibi@on of gene expression by molecules caused by small interfering s (sis) sis and mis similar but form from different precursors sis play a role in heterochroma@n forma@on by facilita@ng methyla@on can block large regions of the chromosome may also block transcrip@on of specific genes 9
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