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ulating dna repair enzymes 80 the effects of mirna dysregulation of gene expression also seem to be important in neuropsychiatric disorders such as schizophrenia bipolar disorder major depression parkinson s disease alzheimer s disease and autism spectrum disorders 81 82 translational edit neomycin is an example of a small molecule that reduces expression of all protein genes inevitably leading to cell death it thus acts as an antibiotic main article translation genetics direct regulation of translation is less prevalent than control of transcription or mrna stability but is occasionally used 83 inhibition of protein translation is a major target for toxins and antibiotics so they can kill a cell by overriding its normal gene expression control 84 protein synthesis inhibitors include the antibiotic neomycin and the toxin ricin 85 post translational modifications edit main article post translational modification post translational modifications ptms are covalent modifications to proteins like rna splicing they help to significantly diversify the proteome these modifications are usually catalyzed by enzymes additionally processes like covalent additions to amino acid side chain residues can often be reversed by other enzymes however some like the proteolytic cleavage of the protein backbone are irreversible 86 ptms play many important roles in the cell 87 for example phosphorylation is primarily involved in activating and deactivating proteins and in signaling pathways 88 ptms are involved in transcriptional regulation an important function of acetylation and methylation is histone tail modification which alters how accessible dna is for transcription 86 they can also be seen in the immune system where glycosylation plays a key role 89 one type of ptm can initiate another type of ptm as can be seen in how ubiquitination tags proteins for degradation through proteolysis 86 proteolysis other than being involved in breaking down proteins is also important in activating and deactivating them and in regulating biological processes such as dna transcription and cell death 90 measurement edit schematic karyogram of a human showing an overview of the expression of the human genome using g banding which is a method that includes giemsa staining wherein the lighter staining regions are generally more transcriptionally active whereas darker regions are more inactive further information karyotype measuring gene expression is an important part of many life sciences as the ability to quantify the level at which a particular gene is expressed within a cell tissue or organism can provide a lot of valuable information for example measuring gene expression can identify viral infection of a cell viral protein expression determine an individual s susceptibility to cancer oncogene expression find if a bacterium is resistant to penicillin beta lactamase expression gene expression profiling evaluates a panel of genes to help understand the fundamental mechanism of a cell this is increasingly used in cancer therapy to target specific chemotherapy see rna seq and dna microarray for details similarly the analysis of the location of protein expression is a powerful tool and this can be done on an organismal or cellular scale investigation of localization is particularly important for the study of development in multicellular organisms and as an indicator of protein function in single cells ideally measurement of expression is done by detecting the final gene product for many genes this is the protein however it is often easier to detect one of the precursors typically mrna and to infer gene expression levels from these measurements mrna quantification edit levels of mrna can be quantitatively measured by northern blotting which provides size and sequence information about the mrna molecules 91 a sample of rna is separated on an agarose gel and hybridized to a radioactively labeled rna probe that is complementary to the target sequence 92 the radiolabeled rna is then detected by an autoradiograph 93 because the use of radioactive reagents makes the procedure time consuming and potentially dangerous alternative labeling and detection methods such as digoxigenin and biotin chemistries have been developed 94 perceived disadvantages of northern blotting are that large quantities of rna are required and that quantification may not be completely accurate as it involves measuring band strength in an image of a gel 95 on the other hand the additional mrna size information from the northern blot allows the discrimination of alternately spliced transcripts 96 97 another approach for measuring mrna abundance is rt qpcr in this technique reverse transcription is followed by quantitative pcr reverse transcription first generates a dna template from the mrna this single stranded template is called cdna the cdna template is then amplified in the quantitative step during which the fluorescence emitted by labeled hybridization probes or intercalating dyes changes as the dna amplification process progresses 98 with a carefully constructed standard curve qpcr can produce an absolute measurement of the number of copies of original mrna typically in units of copies per nanolitre of homogenized tissue or copies per cell 99 qpcr is very sensitive detection of a single mrna molecule is theoretically possible but can be expensive depending on the type of reporter used fluorescently labeled oligonucleotide probes are more expensive than non specific intercalating fluorescent dyes 100 for expression profiling or high throughput analysis of many genes within a sample quantitative pcr may be performed for hundreds of genes simultaneously in the case of low density arrays 101 a second approach is the hybridization microarray a single array or chip may contain probes to determine transcript levels for every known gene in the genome of one or more organisms 102 alternatively tag based technologies like serial analysis of gene expression sage and rna seq which can provide a relative measure of the cellular concentration of different mrnas can be used 103 an advantage of tag based methods is the open architecture allowing for the exact measurement of any transcript with a known or unknown sequence 104 next generation sequencing ngs such as rna seq is another approach producing vast quantities of sequence data that can be matched to a reference genome although ngs is comparatively time consuming expensive and resource intensive it can identify single nucleotide polymorphisms splice variants and novel genes and can also be used to profile expression in organisms for which little or no sequence information is available 105 protein quantification edit for genes encoding proteins the expression level can be directly assessed by a number of methods with some clear analogies to the techniques for mrna quantification one of the most commonly used methods is to perform a western blot against the protein of interest 106 this gives information on the size of the protein in addition to its identity a sample often cellular lysate is separated on a polyacrylamide gel transferred to a membrane and then probed with an antibody to the protein of interest the antibody can either be conjugated to a fluorophore or to horseradish peroxidase for imaging and or quantification the gel based nature of this assay makes quantification less accurate but it has the advantage of being able to identify later modifications to the protein for example proteolysis or ubiquitination from changes in size mrna protein correlation edit while transcription directly reflects gene expression the copy number of mrna molecules does not directly correlate with the number of protein molecules translated from mrna quantification of both protein and mrna permits a correlation of the two levels regulation on each step of gene expression can impact the correlation as shown for regulation of translation 27 or protein stability 107 post translational factors such as protein transport in highly polar cells 108 can influence the measured mrna protein correlation as well localization edit main articles in situ hybridization and immunofluorescence in situ hybridization of drosophila embryos at different developmental stages for the mrna responsible for the expression of hunchback high intensity of blue color marks places with high hunchback mrna quantity analysis of expression is not limited to quantification localization can also be determined mrna can be detected with a suitably labelled complementary mrna strand and protein can be detected via labelled antibodies the probed sample is then observed by microscopy to identify where the mrna or protein is the three dimensional structure of green fluorescent protein the residues in the centre of the barrel are responsible for production of green light after exposing to higher energetic blue light from pdb 1ema by replacing the gene with a new version fused to a green fluorescent protein marker or similar expression may be directly quantified in live cells this is done by imaging using a fluorescence microscope it is very difficult to clone a gfp fused protein into its native location in the genome without affecting expression levels so this method often cannot be used to measure endogenous gene expression it is however widely used to measure the expression of a gene artificially introduced into the cell for example via an expression vector by fusing a target protein to a fluorescent reporter the protein s behavior including its cellular localization and expression level can be significantly changed the enzyme linked immunosorbent assay works by using antibodies immobilised on a microtiter plate to capture proteins of interest from samples added to the well using a detection antibody conjugated to an enzyme or fluorophore the quantity of bound protein can be accurately measured by fluorometric or colourimetric detection the detection process is very similar to that of a western blot but by avoiding the gel steps more accurate quantification can be achieved see also edit allomap molecular expression testing bookmarking epic seq expressed sequence tag expression atlas expression profiling gene structure genetic engineering genetically modified organism list of biological databases list of human genes oscillating gene paramutation protein production protein purification ribonomics ridge sequence profiling tool transcriptional bursting transcriptional noise transcript of unknown function references edit brody l gene expression national human genome research institute nhgri brueckner f armache kj cheung a damsma ge kettenberger h lehmann e et al february 2009 structure function studies of the rna polymerase ii elongation complex acta crystallographica section d biological crystallography 65 pt 2 112 120 bibcode 2009accrd 65 112b doi 10 1107 s0907444908039875 pmc 2631633 pmid 19171965 krebs je goldstein es kilpatrick st 2017 03 02 lewin 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