Allometry of

The cell surface area (SA) increase with volume (V) is determined by growth and regulation of size and shape. Most studies of the rod-shaped model bacteriumEscherichia colihave focussed on the phenomenology or molecular mechanisms governing such scaling. Here, we proceed to examine the role of population statistics and cell division dynamics in such scaling by a combination of microscopy, image analysis and statistical simulations. We find that while the SA of cells sampled from mid-log cultures scales with V by a scaling exponent 2/3, i.e. the geometric law SA ∼V2/3, filamentous cells have higher exponent values. We modulate the growth rate to change the proportion of filamentous cells, and find SA-V scales with an exponent>2/3, exceeding that predicted by the geometric scaling law. However, since increasing growth rates alter the mean and spread of population cell size distributions, we use statistical modeling to disambiguate between the effect of the mean size and variability. Simulating (i) increasing mean cell length with a constant standard deviation (s.d.), (ii) a constant mean length with increasing s.d. and (iii) varying both simultaneously, results in scaling exponents that exceed the 2/3 geometric law, when population variability is included, with the s.d. having a stronger effect. In order to overcome possible effects of statistical sampling of unsynchronized cell populations, we 'virtually synchronized' time-series of cells by using the frames between birth and division identified by the image-analysis pipeline and divided them into four equally spaced phases-B, C1, C2 and D. Phase-specific scaling exponents estimated from these time series and the cell length variability were both found to decrease with the successive stages of birth (B), C1, C2 and division (D). These results point to a need to consider population statistics and a role for cell growth and division when estimating SA-V scaling of bacterial cells.