The messenger RNA has steady state . Since biological translation produces proteins at rate , setting the protein degradation rate equal to this production rate gives
The translational burst size is the expected number of proteins made during one messenger RNA lifetime, whose expected value is . It is dimensionless, rather than a production rate. With independent Poisson process translation and an exponential distribution for the lifetime, the resulting count has a geometric distribution on nonnegative integers with expected value and variance .
With zero initial messenger RNA and protein concentration, solving the two linear ordinary differential equations in sequence gives
The last expression applies after the short messenger RNA transient when . All three mean curves have this shape. At a fixed steady state, a larger translational burst size requires fewer transcription events: . Thus gives infrequent large upward jumps followed by protein degradation, whereas gives many smaller jumps and a smoother trace.
Noise depends on molecule counts. If the fixed cell volume is , write . In the instantaneous-burst limit, independent Poisson process arrivals of geometrically distributed bursts, with count arrival rate , and unit protein degradation give
Indeed, the moment equations are and . With zero initial counts the variance becomes . These equations explain both the approach to the steady state and the scatter about it.
For illustration only, gives at the higher concentration: the two values of the coefficient of variation are and . At the lower concentration, and gives . Decreasing the concentration by a factor of 100 therefore increases relative fluctuations tenfold, despite keeping small bursts. Absolute fluctuation sizes cannot be inferred without a cell volume. The original sketches below stipulate that volume; they are model realizations, not source data.[ref]
Figure 1.
Translational burst size, relative expression fluctuations, and paired fluorescent reporters
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The protein production rate is proportional to the current messenger RNA concentration, not an independently constant : . Setting the two coupled rate equations to zero gives
With first-order removal, a messenger RNA molecule has mean lifetime . At biological translation rate per molecule, it produces an expected proteins before its removal. Thus the translational burst size is a dimensionless mean number of offspring proteins per messenger RNA, not a concentration and not a fixed number produced in every event. When production and removal are independent Poisson events, mixing a Poisson number of biological translation events over an exponentially distributed lifetime makes the burst count geometrically distributed on the nonnegative integers. Its mean is and its variance is .
The protein fluctuation equation gives . With the stated independent noises,
Partial fractions and the supplied integral give the general variance
The specific count-noise formula in the question additionally uses the usual steady birth-and-death noise strengths and . Then reduces it to . Arbitrary unspecified do not by themselves imply this particular Fano factor.
The requested rewriting is
Here is the translational burst size, the expected number of proteins produced during one RNA's mean lifetime. The ratio compares removal rates, equivalently RNA to protein mean lifetimes. Poisson molecule-number statistics have variance equal to mean, hence .
When RNA is much less stable than protein, and . At fixed degradation rates it rises linearly with the rate of biological translation, with exact slope . The formal intercept at zero rate is the limit from positive protein mean; at exactly zero production both mean and variance vanish and their ratio is undefined. A larger Fano factor need not imply a larger relative variance , because the mean also increases.
Figure 1.
Protein Fano factor versus dimensionless rate of biological translation for a short-lived RNA and stable protein: exact two-stage result, burst approximation, and the Poisson baseline
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