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A primary cause of the variability (noise) in gene expression occurring between cells in isogenic populations ^[1]. A major source of transcriptional noise is likely to be transcriptional bursting^[2]^[3]^[4]. Bursting transcription, as opposed to simple probabilistic models of transcription, reflects multiple states of gene activity, with fluctuations between states separated by irregular intervals, generating uneven protein expression between cells. Noise in gene expression can have tremendous consequences on cell behaviour, and must be mitigated or integrated. In certain contexts, such as the survival of microbes in rapidly changing stressful environments, or several types of scattered differentiation, the variability may be essential. Variability also impacts upon the effectiveness of clinical treatment, with resistance of bacteria to antibiotics demonstrably caused by non-genetic differences^[5]. Variability in gene expression may also contribute to resistance of sub-populations of cancer cells to chemotherapy^[6].

Notes

^ Raj, A; Van Oudenaarden, A (2008). "Nature, nurture, or chance: stochastic gene expression and its consequences". Cell. 135 (2): 216–26. doi:10.1016/j.cell.2008.09.050. PMID 18957198.
^ Golding, I; Paulsson, J; Zawilski, SM; Cox, EC (2005). "Real-time kinetics of gene activity in individual bacteria". Cell. 123 (6): 1025–36. doi:10.1016/j.cell.2005.09.031. PMID 16360033.
^ Chubb, JR; Trcek, T; Shenoy, SM; Singer, RH (2006). "Transcriptional pulsing of a developmental gene". Current biology : CB. 16 (10): 1018–25. doi:10.1016/j.cub.2006.03.092. PMID 16713960.
^ Raj, A; Peskin, CS; Tranchina, D; Vargas, DY; Tyagi, S (2006). "Stochastic mRNA synthesis in mammalian cells". PLoS biology. 4 (10): e309. doi:10.1371/journal.pbio.0040309. PMC 1563489. PMID 17048983.{{cite journal}}: CS1 maint: unflagged free DOI (link)
^ Attention: This template ({{cite pmid}}) is deprecated. To cite the publication identified by PMID 18388284, please use {{cite journal}} with |pmid=18388284 instead.
^ Attention: This template ({{cite pmid}}) is deprecated. To cite the publication identified by PMID 6348026, please use {{cite journal}} with |pmid=6348026 instead.

[1] Raj, A; Van Oudenaarden, A (2008). "Nature, nurture, or chance: stochastic gene expression and its consequences". Cell. 135 (2): 216–26. doi:10.1016/j.cell.2008.09.050. PMID 18957198.

[2] Golding, I; Paulsson, J; Zawilski, SM; Cox, EC (2005). "Real-time kinetics of gene activity in individual bacteria". Cell. 123 (6): 1025–36. doi:10.1016/j.cell.2005.09.031. PMID 16360033.

[3] Chubb, JR; Trcek, T; Shenoy, SM; Singer, RH (2006). "Transcriptional pulsing of a developmental gene". Current biology : CB. 16 (10): 1018–25. doi:10.1016/j.cub.2006.03.092. PMID 16713960.

[4] Raj, A; Peskin, CS; Tranchina, D; Vargas, DY; Tyagi, S (2006). "Stochastic mRNA synthesis in mammalian cells". PLoS biology. 4 (10): e309. doi:10.1371/journal.pbio.0040309. PMC 1563489. PMID 17048983.{{cite journal}}: CS1 maint: unflagged free DOI (link)

[5] Attention: This template ({{cite pmid}}) is deprecated. To cite the publication identified by PMID 18388284, please use {{cite journal}} with |pmid=18388284 instead.

[6] Attention: This template ({{cite pmid}}) is deprecated. To cite the publication identified by PMID 6348026, please use {{cite journal}} with |pmid=6348026 instead.

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	A primary cause of the variability ([[noise]]) in [[gene expression]] occurring between cells in isogenic populations <ref>{{cite journal \|pmid=18957198 \|year=2008 \|last1=Raj \|first1=A \|last2=Van Oudenaarden \|first2=A \|title=Nature, nurture, or chance: stochastic gene expression and its consequences \|volume=135 \|issue=2 \|pages=216–26 \|doi=10.1016/j.cell.2008.09.050 \|journal=Cell}}</ref>. A major source of transcriptional noise is likely to be [[transcriptional bursting]]<ref>{{cite journal \|pmid=16360033 \|year=2005 \|last1=Golding \|first1=I \|last2=Paulsson \|first2=J \|last3=Zawilski \|first3=SM \|last4=Cox \|first4=EC \|title=Real-time kinetics of gene activity in individual bacteria \|volume=123 \|issue=6 \|pages=1025–36 \|doi=10.1016/j.cell.2005.09.031 \|journal=Cell}}</ref><ref>{{cite journal \|pmid=16713960 \|year=2006 \|last1=Chubb \|first1=JR \|last2=Trcek \|first2=T \|last3=Shenoy \|first3=SM \|last4=Singer \|first4=RH \|title=Transcriptional pulsing of a developmental gene \|volume=16 \|issue=10 \|pages=1018–25 \|doi=10.1016/j.cub.2006.03.092 \|journal=Current biology : CB}}</ref><ref>{{cite journal \|pmid=17048983 \|year=2006 \|last1=Raj \|first1=A \|last2=Peskin \|first2=CS \|last3=Tranchina \|first3=D \|last4=Vargas \|first4=DY \|last5=Tyagi \|first5=S \|title=Stochastic mRNA synthesis in mammalian cells \|volume=4 \|issue=10 \|pages=e309 \|doi=10.1371/journal.pbio.0040309 \|pmc=1563489 \|journal=PLoS biology}}</ref>. Bursting transcription, as opposed to simple probabilistic models of transcription, reflects multiple states of gene activity, with fluctuations between states separated by irregular intervals, generating uneven protein expression between cells <ref>{{cite journal \|pmid=18957198 \|year=2008 \|last1=Raj \|first1=A \|last2=Van Oudenaarden \|first2=A \|title=Nature, nurture, or chance: stochastic gene expression and its consequences \|volume=135 \|issue=2 \|pages=216–26 \|doi=10.1016/j.cell.2008.09.050 \|journal=Cell}}</ref>. Noise in gene expression can have tremendous consequences on cell behaviour, and must be mitigated or integrated. In certain contexts, such as the survival of microbes in rapidly changing stressful environments, or several types of scattered differentiation, the variability may be essential. Variability also impacts upon the effectiveness of clinical treatment, with resistance of bacteria to [[antibiotics]] demonstrably caused by non-genetic differences<ref>{{cite pmid\|~~19337290~~}}</ref>. Variability in gene expression may also contribute to resistance of sub-populations of cancer cells to chemotherapy<ref>{{cite pmid\|6348026}}</ref>.		A primary cause of the variability ([[noise]]) in [[gene expression]] occurring between cells in isogenic populations <ref>{{cite journal \|pmid=18957198 \|year=2008 \|last1=Raj \|first1=A \|last2=Van Oudenaarden \|first2=A \|title=Nature, nurture, or chance: stochastic gene expression and its consequences \|volume=135 \|issue=2 \|pages=216–26 \|doi=10.1016/j.cell.2008.09.050 \|journal=Cell}}</ref>. A major source of transcriptional noise is likely to be [[transcriptional bursting]]<ref>{{cite journal \|pmid=16360033 \|year=2005 \|last1=Golding \|first1=I \|last2=Paulsson \|first2=J \|last3=Zawilski \|first3=SM \|last4=Cox \|first4=EC \|title=Real-time kinetics of gene activity in individual bacteria \|volume=123 \|issue=6 \|pages=1025–36 \|doi=10.1016/j.cell.2005.09.031 \|journal=Cell}}</ref><ref>{{cite journal \|pmid=16713960 \|year=2006 \|last1=Chubb \|first1=JR \|last2=Trcek \|first2=T \|last3=Shenoy \|first3=SM \|last4=Singer \|first4=RH \|title=Transcriptional pulsing of a developmental gene \|volume=16 \|issue=10 \|pages=1018–25 \|doi=10.1016/j.cub.2006.03.092 \|journal=Current biology : CB}}</ref><ref>{{cite journal \|pmid=17048983 \|year=2006 \|last1=Raj \|first1=A \|last2=Peskin \|first2=CS \|last3=Tranchina \|first3=D \|last4=Vargas \|first4=DY \|last5=Tyagi \|first5=S \|title=Stochastic mRNA synthesis in mammalian cells \|volume=4 \|issue=10 \|pages=e309 \|doi=10.1371/journal.pbio.0040309 \|pmc=1563489 \|journal=PLoS biology}}</ref>. Bursting transcription, as opposed to simple probabilistic models of transcription, reflects multiple states of gene activity, with fluctuations between states separated by irregular intervals, generating uneven protein expression between cells. Noise in gene expression can have tremendous consequences on cell behaviour, and must be mitigated or integrated. In certain contexts, such as the survival of microbes in rapidly changing stressful environments, or several types of scattered differentiation, the variability may be essential. Variability also impacts upon the effectiveness of clinical treatment, with resistance of bacteria to [[antibiotics]] demonstrably caused by non-genetic differences<ref>{{cite pmid\|18388284}}</ref>. Variability in gene expression may also contribute to resistance of sub-populations of cancer cells to chemotherapy<ref>{{cite pmid\|6348026}}</ref>.