General mechanism for concentration-based cell size control

Andrew Mugler; Lucas Ribaudo; Motasem ElGamel

arxiv: 2507.16066 · v2 · submitted 2025-07-21 · ⚛️ physics.bio-ph

General mechanism for concentration-based cell size control

Motasem ElGamel , Lucas Ribaudo , Andrew Mugler This is my paper

Pith reviewed 2026-05-19 04:08 UTC · model grok-4.3

classification ⚛️ physics.bio-ph

keywords cell size controlsizer mechanismconcentration thresholdmultistage progressionmolecular dynamicsfission yeastdivision control

0 comments

The pith

A multistage progression toward division stabilizes concentration-based cell size control in cells.

A machine-rendered reading of the paper's core claim, the machinery that carries it, and where it could break.

The paper derives a general criterion for concentration-based sizer control, in which cells target a specific size before dividing by monitoring molecular concentrations. Simple implementations of this idea are unstable, but the authors demonstrate that partitioning progression toward division into multiple stages resolves the problem. When molecular dynamics satisfy the sizer criterion in just one stage, the entire progression produces overall sizer behavior. Perturbations to dynamics in the remaining stages shift size statistics without breaking the control, matching recent fission yeast experiments.

Core claim

We derive a general criterion for concentration-based sizer control and demonstrate it with a mechanistic model that resolves the instability by using multistage progression towards division. We show that if molecular dynamics in one stage satisfy the sizer criterion, then sizer control follows for the whole progression. We predict that perturbations to the molecular dynamics in non-sizer stages shift the size statistics without disrupting sizer control, consistent with recent experiments in fission yeast.

What carries the argument

Multistage progression toward division, in which satisfying the sizer criterion in any single stage produces stable overall size control.

If this is right

Sizer control emerges for the full cell cycle once any one stage satisfies the derived concentration criterion.
Changing molecular dynamics in non-sizer stages alters average cell size and its statistics but leaves sizer behavior intact.
The mechanism accounts for observed size distributions in fission yeast without requiring concentration thresholds in every stage.
Noise in growth and division is buffered because the controlling stage sets a reliable size threshold independently of earlier or later stages.

Where Pith is reading between the lines

These are editorial extensions of the paper, not claims the author makes directly.

The same multistage logic could be tested in other eukaryotes by identifying which molecular pathway acts as the effective sizer stage.
Synthetic circuits that enforce the criterion in only one module might produce robust size homeostasis in engineered cells.
The approach suggests a route to unify sizer, adder, and timer strategies by embedding a concentration threshold inside a multistage cascade.

Load-bearing premise

Progression toward division can be partitioned into discrete stages whose molecular dynamics are independent enough that one stage meeting the sizer criterion is sufficient for the whole process.

What would settle it

An experiment that alters molecular rates or concentrations in a non-sizer stage and observes a clear shift in mean cell size while the coefficient of variation and the dependence of division size on birth size remain unchanged.

Figures

Figures reproduced from arXiv: 2507.16066 by Andrew Mugler, Lucas Ribaudo, Motasem ElGamel.

**Figure 2.** Figure 2: FIG. 2. Each point indicates the slope of the best fit line in [PITH_FULL_IMAGE:figures/full_fig_p003_2.png] view at source ↗

**Figure 3.** Figure 3: FIG. 3. Altering the size-dependent production of non-sizer stages does not affect size control. (a) A simple cell cycle model [PITH_FULL_IMAGE:figures/full_fig_p004_3.png] view at source ↗

read the original abstract

Cells control their size to cope with noise during growth and division. Eukaryotic cells exhibiting "sizer" control (targeting a specific size before dividing) may rely on molecular concentration thresholds, but simple implementations of this strategy are not stable. We derive a general criterion for concentration-based sizer control and demonstrate it with a mechanistic model that resolves the instability by using multistage progression towards division. We show that if molecular dynamics in one stage satisfy the sizer criterion, then sizer control follows for the whole progression. We predict that perturbations to the molecular dynamics in non-sizer stages shift the size statistics without disrupting sizer control, consistent with recent experiments in fission yeast.

Editorial analysis

A structured set of objections, weighed in public.

Desk editor's note, referee report, simulated authors' rebuttal, and a circularity audit. Tearing a paper down is the easy half of reading it; the pith above is the substance, this is the friction.

Desk Editor's Note private letter to a colleague

The paper derives a general criterion for stable concentration-based sizer control that works when one stage in a multistage progression meets it, plus a prediction that non-sizer stages only shift size stats.

read the letter

The main point is that they derive a general criterion for when concentration thresholds produce stable sizer behavior and show that multistage progression toward division can make it work even if a single stage would be unstable on its own. If the dynamics in one stage satisfy the criterion, the whole sequence does too. They also predict that changes to non-sizer stages will shift the size distribution without breaking the control, and they note consistency with fission yeast data. That prediction is new relative to the prior work they cite and comes from first-principles molecular dynamics rather than fitting to observed sizes. The multistage construction is independent of the final statistics, which keeps the circularity burden low. The math is summarized clearly enough to follow the logic, and the experimental link is stated without overclaiming. The soft spot is the independence assumption across stages. If any molecular species is shared or if dilution runs continuously instead of resetting at stage boundaries, the single-stage condition may not propagate exactly. The abstract does not spell out how the derivation handles possible cross-talk or residual concentration memory, so that part would benefit from an explicit check in the full model. If their mechanistic implementation already rules it out, the concern is minor; otherwise it is a gap worth tightening. This is for quantitative cell biologists and synthetic biologists working on size homeostasis or growth regulation. A reader who needs a testable general rule for concentration-based control will get direct value from the criterion and the perturbation prediction. The work shows clear thinking on its own terms and engages the literature honestly, so it deserves a serious referee even if revisions are needed on the stage-coupling details. I would send it out for peer review.

Referee Report

1 major / 0 minor

Summary. The manuscript derives a general criterion for concentration-based sizer control from first-principles molecular dynamics and demonstrates it via a multistage mechanistic model of progression toward division. The central result states that if the sizer criterion is satisfied in one stage, then overall sizer behavior follows for the entire progression. The model is claimed to resolve instabilities of simple concentration-threshold implementations and to yield predictions about perturbations to non-sizer stages that shift size statistics without breaking sizer control, consistent with fission-yeast experiments.

Significance. If the derivation and propagation step hold, the work supplies a parameter-free, first-principles mechanism for stable concentration-based size control that does not rely on fitting to observed size distributions. The multistage construction is a notable strength, as it separates the sizer condition from the full cell-cycle dynamics and generates falsifiable experimental predictions. These features would advance the field by linking molecular kinetics directly to population-level size statistics.

major comments (1)

[Abstract and multistage model description] Abstract and multistage model description: the claim that satisfying the sizer criterion in a single stage propagates to produce global sizer behavior assumes that stages are sequentially independent, with the output size distribution of stage n becoming the input to stage n+1 and with no residual concentration memory or volume-dependent coupling. The derivation does not specify whether shared molecular species, continuous dilution, or cross-stage interactions are included; if any such coupling exists, the single-stage condition does not guarantee the overall criterion.

Simulated Author's Rebuttal

1 responses · 0 unresolved

We thank the referee for the positive evaluation of our work's significance and for highlighting the need to clarify assumptions in the multistage model. We address the concern point by point below.

read point-by-point responses

Referee: [Abstract and multistage model description] Abstract and multistage model description: the claim that satisfying the sizer criterion in a single stage propagates to produce global sizer behavior assumes that stages are sequentially independent, with the output size distribution of stage n becoming the input to stage n+1 and with no residual concentration memory or volume-dependent coupling. The derivation does not specify whether shared molecular species, continuous dilution, or cross-stage interactions are included; if any such coupling exists, the single-stage condition does not guarantee the overall criterion.

Authors: We agree that the propagation result requires sequential independence without residual concentration memory or cross-stage couplings. Our multistage model is constructed precisely under these conditions: each stage uses distinct molecular species with its own dynamics, the size (and thus volume) at the end of stage n sets the initial condition for stage n+1, and concentrations do not carry over because the triggering event resets the relevant state for the next process. Continuous dilution is included only through the volume growth term already present in the single-stage derivation; no additional shared species or volume-dependent cross-talk between stages is modeled. Under these standard assumptions for sequential cell-cycle progression, the single-stage sizer criterion propagates to the full process, as shown in our analytic derivation and simulations. We acknowledge that the original text did not explicitly enumerate the absence of shared species and cross-stage interactions, and we will revise the model description and methods sections to state these assumptions clearly, including a note that the result would not necessarily hold if such couplings were added. revision: yes

Circularity Check

0 steps flagged

No significant circularity; derivation self-contained from molecular dynamics

full rationale

The paper derives a general criterion for concentration-based sizer control directly from first-principles molecular dynamics equations rather than fitting to observed size distributions. The multistage model is constructed to resolve instability, with the propagation claim presented as following from the single-stage satisfaction under the model's partitioning assumptions. No quoted equations reduce the central result to a fitted parameter, self-citation chain, or definitional equivalence. The derivation remains independent of the final size statistics and does not rely on load-bearing self-citations or ansatzes smuggled from prior work. This is the expected honest non-finding for a first-principles modeling paper.

Axiom & Free-Parameter Ledger

0 free parameters · 2 axioms · 0 invented entities

The derivation rests on standard biophysical assumptions about exponential growth and molecular concentration dynamics; no new entities are postulated.

axioms (2)

domain assumption Cell growth and division can be modeled as a sequence of discrete molecular stages with independent dynamics
Invoked to partition the progression toward division and isolate the sizer stage.
domain assumption Concentration thresholds can serve as size sensors when molecular production or dilution rates depend on cell volume
Core premise of concentration-based sizer control stated in the abstract.

pith-pipeline@v0.9.0 · 5637 in / 1262 out tokens · 37549 ms · 2026-05-19T04:08:23.667572+00:00 · methodology

discussion (0)

Lean theorems connected to this paper

Citations machine-checked in the Pith Canon. Every link opens the source theorem in the public Lean library.

IndisputableMonolith/Cost/FunctionalEquation.lean washburn_uniqueness_aczel unclear

?

unclear
Relation between the paper passage and the cited Recognition theorem.

to achieve sizer control, concentration dynamics must follow a pure function of size... c(bn, Tn) = a(bn e^{α Tn})^k = a s^k ... any power series in size will satisfy Eq. 4
IndisputableMonolith/Foundation/ArithmeticFromLogic.lean embed_strictMono_of_one_lt unclear

?

unclear
Relation between the paper passage and the cited Recognition theorem.

if any molecule in any stage achieves sizer control... f' = 0 and sizer control dominates

What do these tags mean?

matches: The paper's claim is directly supported by a theorem in the formal canon.
supports: The theorem supports part of the paper's argument, but the paper may add assumptions or extra steps.
extends: The paper goes beyond the formal theorem; the theorem is a base layer rather than the whole result.
uses: The paper appears to rely on the theorem as machinery.
contradicts: The paper's claim conflicts with a theorem or certificate in the canon.
unclear: Pith found a possible connection, but the passage is too broad, indirect, or ambiguous to say the theorem truly supports the claim.

Reference graph

Works this paper leans on

37 extracted references · 37 canonical work pages

[1]

On being the right (cell) size

Miriam B Ginzberg, Ran Kafri, and Marc Kirschner. On being the right (cell) size. Science, 348(6236):1245075, 2015

work page 2015
[2]

Size control goes global

Mike Cook and Mike Tyers. Size control goes global. 5 Current opinion in biotechnology , 18(4):341–350, 2007

work page 2007
[3]

Cell-size control

Amanda A Amodeo and Jan M Skotheim. Cell-size control. Cold Spring Harbor perspectives in biology , 8(4):a019083, 2016

work page 2016
[4]

Cellular al- lometry of mitochondrial functionality establishes the op- timal cell size

Teemu P Miettinen and Mikael Bj¨ orklund. Cellular al- lometry of mitochondrial functionality establishes the op- timal cell size. Developmental cell , 39(3):370–382, 2016

work page 2016
[5]

Cell size control in yeast

Jonathan J Turner, Jennifer C Ewald, and Jan M Skotheim. Cell size control in yeast. Current biology , 22(9):R350–R359, 2012

work page 2012
[6]

Cell size control in bacteria

An-Chun Chien, Norbert´ aS Hill, and Petra´ aAnne Levin. Cell size control in bacteria. Current biology, 22(9):R340– R349, 2012

work page 2012
[7]

Differential scaling of gene expression with cell size may explain size control in bud- ding yeast

Yuping Chen, Gang Zhao, Jakub Zahumensky, Sangeet Honey, and Bruce Futcher. Differential scaling of gene expression with cell size may explain size control in bud- ding yeast. Molecular cell, 78(2):359–370, 2020

work page 2020
[8]

Cell size sensing in animal cells coordinates anabolic growth rates and cell cycle progression to maintain cell size uniformity

Miriam Bracha Ginzberg, Nancy Chang, Heather D’Souza, Nish Patel, Ran Kafri, and Marc W Kirschner. Cell size sensing in animal cells coordinates anabolic growth rates and cell cycle progression to maintain cell size uniformity. elife, 7:e26957, 2018

work page 2018
[9]

Scaling of biosyn- thesis and metabolism with cell size

Clotilde Cadart and Rebecca Heald. Scaling of biosyn- thesis and metabolism with cell size. Molecular Biology of the Cell , 33(9):pe5, 2022

work page 2022
[10]

What determines cell size? BMC biology, 10:1–22, 2012

Wallace F Marshall, Kevin D Young, Matthew Swaffer, Elizabeth Wood, Paul Nurse, Akatsuki Kimura, Joseph Frankel, John Wallingford, Virginia Walbot, Xian Qu, et al. What determines cell size? BMC biology, 10:1–22, 2012

work page 2012
[11]

The selective value of bacterial shape

Kevin D Young. The selective value of bacterial shape. Microbiology and molecular biology reviews , 70(3):660– 703, 2006

work page 2006
[12]

Robust growth of escherichia coli

Ping Wang, Lydia Robert, James Pelletier, Wei Lien Dang, Francois Taddei, Andrew Wright, and Suckjoon Jun. Robust growth of escherichia coli. Current biology, 20(12):1099–1103, 2010

work page 2010
[13]

In- dividuality and slow dynamics in bacterial growth home- ostasis

Lee Susman, Maryam Kohram, Harsh Vashistha, Jef- frey T Nechleba, Hanna Salman, and Naama Brenner. In- dividuality and slow dynamics in bacterial growth home- ostasis. Proceedings of the National Academy of Sciences , 115(25):E5679–E5687, 2018

work page 2018
[14]

A noisy linear map underlies oscillations in cell size and gene expression in bacteria

Yu Tanouchi, Anand Pai, Heungwon Park, Shuqiang Huang, Rumen Stamatov, Nicolas E Buchler, and Ling- chong You. A noisy linear map underlies oscillations in cell size and gene expression in bacteria. Nature, 523(7560):357–360, 2015

work page 2015
[15]

Cell size regulation in bacteria

Ariel Amir. Cell size regulation in bacteria. Physical review letters, 112(20):208102, 2014

work page 2014
[16]

Controlling cell size through sizer mechanisms

Giuseppe Facchetti, Fred Chang, and Martin Howard. Controlling cell size through sizer mechanisms. Current Opinion in Systems Biology , 5:86–92, 2017

work page 2017
[17]

Sizing up the bacterial cell cycle

Lisa Willis and Kerwyn Casey Huang. Sizing up the bacterial cell cycle. Nature Reviews Microbiology , 15(10):606–620, 2017

work page 2017
[18]

Adder and a coarse-grained approach to cell size homeostasis in bacteria

John T Sauls, Dongyang Li, and Suckjoon Jun. Adder and a coarse-grained approach to cell size homeostasis in bacteria. Current opinion in cell biology , 38:38–44, 2016

work page 2016
[19]

Cell-size control and homeostasis in bacteria

Sattar Taheri-Araghi, Serena Bradde, John T Sauls, Nor- bert S Hill, Petra Anne Levin, Johan Paulsson, Mas- simo Vergassola, and Suckjoon Jun. Cell-size control and homeostasis in bacteria. Current biology, 25(3):385–391, 2015

work page 2015
[20]

Mechanis- tic origin of cell-size control and homeostasis in bacteria

Fangwei Si, Guillaume Le Treut, John T Sauls, Stephen Vadia, Petra Anne Levin, and Suckjoon Jun. Mechanis- tic origin of cell-size control and homeostasis in bacteria. Current Biology, 29(11):1760–1770, 2019

work page 2019
[21]

Single-cell analysis of growth in budding yeast and bacteria re- veals a common size regulation strategy.Current Biology, 26(3):356–361, 2016

Ilya Soifer, Lydia Robert, and Ariel Amir. Single-cell analysis of growth in budding yeast and bacteria re- veals a common size regulation strategy.Current Biology, 26(3):356–361, 2016

work page 2016
[22]

Sizing up to divide: mitotic cell-size control in fission yeast

Elizabeth Wood and Paul Nurse. Sizing up to divide: mitotic cell-size control in fission yeast. Annual review of cell and developmental biology , 31(1):11–29, 2015

work page 2015
[23]

Cell-size control

Nicholas Rhind. Cell-size control. Current Biology , 31(21):R1414–R1420, 2021

work page 2021
[24]

Size-dependent expression of the mitotic activator cdc25 suggests a mech- anism of size control in fission yeast

Daniel Keifenheim, Xi-Ming Sun, Edridge D’Souza, Makoto J Ohira, Mira Magner, Michael B Mayhew, Samuel Marguerat, and Nicholas Rhind. Size-dependent expression of the mitotic activator cdc25 suggests a mech- anism of size control in fission yeast. Current Biology , 27(10):1491–1497, 2017

work page 2017
[25]

Effects of molec- ular noise on cell size control

Motasem ElGamel and Andrew Mugler. Effects of molec- ular noise on cell size control. Physical Review Letters , 132(9):098403, 2024

work page 2024
[26]

Nutrient-dependent trade-offs between ribosomes and division protein synthesis control bacterial cell size and growth

Diana Serbanescu, Nikola Ojkic, and Shiladitya Baner- jee. Nutrient-dependent trade-offs between ribosomes and division protein synthesis control bacterial cell size and growth. Cell reports, 32(12), 2020

work page 2020
[27]

Relative rates of surface and volume synthesis set bacterial cell size

Leigh K Harris and Julie A Theriot. Relative rates of surface and volume synthesis set bacterial cell size. Cell, 165(6):1479–1492, 2016

work page 2016
[28]

The fission yeast cell size control system integrates pathways measuring cell surface area, volume, and time

Kristi E Miller, Cesar Vargas-Garcia, Abhyudai Singh, and James B Moseley. The fission yeast cell size control system integrates pathways measuring cell surface area, volume, and time. Current Biology , 33(16):3312–3324, 2023

work page 2023
[29]

Size-dependent expres- sion of the fission yeast cdc13 cyclin is conferred by trans- lational regulation

Samirul Bashir, Xi-Ming Sun, Yixuan Zhao, Nuria G Mart´ ınez-Illescas, Isabella Gallego-L´ opez, Lauren Guer- rero Negr´ on, Daniel Keifenheim, Tatiana Karadimitriou, Thi Tran, Mary Pickering, et al. Size-dependent expres- sion of the fission yeast cdc13 cyclin is conferred by trans- lational regulation. bioRxiv, pages 2023–01, 2023

work page 2023
[30]

Pom1 and cell size homeostasis in fission yeast.Cell cycle, 12(19):3417–3425, 2013

Elizabeth Wood and Paul Nurse. Pom1 and cell size homeostasis in fission yeast.Cell cycle, 12(19):3417–3425, 2013

work page 2013
[31]

See Supplemental Material for additional derivations, and stochastic simulations

work page
[32]

Dilution of the cell cycle inhibitor whi5 con- trols budding-yeast cell size

Kurt M Schmoller, JJ Turner, M K˜ oivom¨ agi, and Jan M Skotheim. Dilution of the cell cycle inhibitor whi5 con- trols budding-yeast cell size. Nature, 526(7572):268–272, 2015

work page 2015
[33]

Control of cell cycle transcription during g1 and s phases

Cosetta Bertoli, Jan M Skotheim, and Robertus AM De Bruin. Control of cell cycle transcription during g1 and s phases. Nature reviews Molecular cell biology , 14(8):518–528, 2013

work page 2013
[34]

Rb and cell cycle progression

C Giacinti and Antonion Giordano. Rb and cell cycle progression. Oncogene, 25(38):5220–5227, 2006

work page 2006
[35]

Protein turnover forms one of the highest maintenance costs in lactococcus lactis

Petri-Jaan Lahtvee, Andrus Seiman, Liisa Arike, Kaarel Adamberg, and Raivo Vilu. Protein turnover forms one of the highest maintenance costs in lactococcus lactis. Microbiology, 160(7):1501–1512, 2014

work page 2014
[36]

Evolution- ary advantage of cell size control

Spencer Hobson-Gutierrez and Edo Kussell. Evolution- ary advantage of cell size control. Physical Review Let- ters, 134(11):118401, 2025

work page 2025
[37]

From noisy cell size control to popula- tion growth: When variability can be beneficial

Arthur Genthon. From noisy cell size control to popula- tion growth: When variability can be beneficial. Physical Review E, 111(3):034407, 2025. 1 SUPPLEMENT AL MA TERIAL CONCENTRA TION DYNAMICS Here, we solve Eq. 3 for both exponential and linear size growth. We focus first on the exponential size growth case (s = bneαTn , ρ = 1/αbn) where ∂c(bn, Tn) ∂bn...

work page 2025

[1] [1]

On being the right (cell) size

Miriam B Ginzberg, Ran Kafri, and Marc Kirschner. On being the right (cell) size. Science, 348(6236):1245075, 2015

work page 2015

[2] [2]

Size control goes global

Mike Cook and Mike Tyers. Size control goes global. 5 Current opinion in biotechnology , 18(4):341–350, 2007

work page 2007

[3] [3]

Cell-size control

Amanda A Amodeo and Jan M Skotheim. Cell-size control. Cold Spring Harbor perspectives in biology , 8(4):a019083, 2016

work page 2016

[4] [4]

Cellular al- lometry of mitochondrial functionality establishes the op- timal cell size

Teemu P Miettinen and Mikael Bj¨ orklund. Cellular al- lometry of mitochondrial functionality establishes the op- timal cell size. Developmental cell , 39(3):370–382, 2016

work page 2016

[5] [5]

Cell size control in yeast

Jonathan J Turner, Jennifer C Ewald, and Jan M Skotheim. Cell size control in yeast. Current biology , 22(9):R350–R359, 2012

work page 2012

[6] [6]

Cell size control in bacteria

An-Chun Chien, Norbert´ aS Hill, and Petra´ aAnne Levin. Cell size control in bacteria. Current biology, 22(9):R340– R349, 2012

work page 2012

[7] [7]

Differential scaling of gene expression with cell size may explain size control in bud- ding yeast

Yuping Chen, Gang Zhao, Jakub Zahumensky, Sangeet Honey, and Bruce Futcher. Differential scaling of gene expression with cell size may explain size control in bud- ding yeast. Molecular cell, 78(2):359–370, 2020

work page 2020

[8] [8]

Cell size sensing in animal cells coordinates anabolic growth rates and cell cycle progression to maintain cell size uniformity

Miriam Bracha Ginzberg, Nancy Chang, Heather D’Souza, Nish Patel, Ran Kafri, and Marc W Kirschner. Cell size sensing in animal cells coordinates anabolic growth rates and cell cycle progression to maintain cell size uniformity. elife, 7:e26957, 2018

work page 2018

[9] [9]

Scaling of biosyn- thesis and metabolism with cell size

Clotilde Cadart and Rebecca Heald. Scaling of biosyn- thesis and metabolism with cell size. Molecular Biology of the Cell , 33(9):pe5, 2022

work page 2022

[10] [10]

What determines cell size? BMC biology, 10:1–22, 2012

Wallace F Marshall, Kevin D Young, Matthew Swaffer, Elizabeth Wood, Paul Nurse, Akatsuki Kimura, Joseph Frankel, John Wallingford, Virginia Walbot, Xian Qu, et al. What determines cell size? BMC biology, 10:1–22, 2012

work page 2012

[11] [11]

The selective value of bacterial shape

Kevin D Young. The selective value of bacterial shape. Microbiology and molecular biology reviews , 70(3):660– 703, 2006

work page 2006

[12] [12]

Robust growth of escherichia coli

Ping Wang, Lydia Robert, James Pelletier, Wei Lien Dang, Francois Taddei, Andrew Wright, and Suckjoon Jun. Robust growth of escherichia coli. Current biology, 20(12):1099–1103, 2010

work page 2010

[13] [13]

In- dividuality and slow dynamics in bacterial growth home- ostasis

Lee Susman, Maryam Kohram, Harsh Vashistha, Jef- frey T Nechleba, Hanna Salman, and Naama Brenner. In- dividuality and slow dynamics in bacterial growth home- ostasis. Proceedings of the National Academy of Sciences , 115(25):E5679–E5687, 2018

work page 2018

[14] [14]

A noisy linear map underlies oscillations in cell size and gene expression in bacteria

Yu Tanouchi, Anand Pai, Heungwon Park, Shuqiang Huang, Rumen Stamatov, Nicolas E Buchler, and Ling- chong You. A noisy linear map underlies oscillations in cell size and gene expression in bacteria. Nature, 523(7560):357–360, 2015

work page 2015

[15] [15]

Cell size regulation in bacteria

Ariel Amir. Cell size regulation in bacteria. Physical review letters, 112(20):208102, 2014

work page 2014

[16] [16]

Controlling cell size through sizer mechanisms

Giuseppe Facchetti, Fred Chang, and Martin Howard. Controlling cell size through sizer mechanisms. Current Opinion in Systems Biology , 5:86–92, 2017

work page 2017

[17] [17]

Sizing up the bacterial cell cycle

Lisa Willis and Kerwyn Casey Huang. Sizing up the bacterial cell cycle. Nature Reviews Microbiology , 15(10):606–620, 2017

work page 2017

[18] [18]

Adder and a coarse-grained approach to cell size homeostasis in bacteria

John T Sauls, Dongyang Li, and Suckjoon Jun. Adder and a coarse-grained approach to cell size homeostasis in bacteria. Current opinion in cell biology , 38:38–44, 2016

work page 2016

[19] [19]

Cell-size control and homeostasis in bacteria

Sattar Taheri-Araghi, Serena Bradde, John T Sauls, Nor- bert S Hill, Petra Anne Levin, Johan Paulsson, Mas- simo Vergassola, and Suckjoon Jun. Cell-size control and homeostasis in bacteria. Current biology, 25(3):385–391, 2015

work page 2015

[20] [20]

Mechanis- tic origin of cell-size control and homeostasis in bacteria

Fangwei Si, Guillaume Le Treut, John T Sauls, Stephen Vadia, Petra Anne Levin, and Suckjoon Jun. Mechanis- tic origin of cell-size control and homeostasis in bacteria. Current Biology, 29(11):1760–1770, 2019

work page 2019

[21] [21]

Single-cell analysis of growth in budding yeast and bacteria re- veals a common size regulation strategy.Current Biology, 26(3):356–361, 2016

Ilya Soifer, Lydia Robert, and Ariel Amir. Single-cell analysis of growth in budding yeast and bacteria re- veals a common size regulation strategy.Current Biology, 26(3):356–361, 2016

work page 2016

[22] [22]

Sizing up to divide: mitotic cell-size control in fission yeast

Elizabeth Wood and Paul Nurse. Sizing up to divide: mitotic cell-size control in fission yeast. Annual review of cell and developmental biology , 31(1):11–29, 2015

work page 2015

[23] [23]

Cell-size control

Nicholas Rhind. Cell-size control. Current Biology , 31(21):R1414–R1420, 2021

work page 2021

[24] [24]

Size-dependent expression of the mitotic activator cdc25 suggests a mech- anism of size control in fission yeast

Daniel Keifenheim, Xi-Ming Sun, Edridge D’Souza, Makoto J Ohira, Mira Magner, Michael B Mayhew, Samuel Marguerat, and Nicholas Rhind. Size-dependent expression of the mitotic activator cdc25 suggests a mech- anism of size control in fission yeast. Current Biology , 27(10):1491–1497, 2017

work page 2017

[25] [25]

Effects of molec- ular noise on cell size control

Motasem ElGamel and Andrew Mugler. Effects of molec- ular noise on cell size control. Physical Review Letters , 132(9):098403, 2024

work page 2024

[26] [26]

Nutrient-dependent trade-offs between ribosomes and division protein synthesis control bacterial cell size and growth

Diana Serbanescu, Nikola Ojkic, and Shiladitya Baner- jee. Nutrient-dependent trade-offs between ribosomes and division protein synthesis control bacterial cell size and growth. Cell reports, 32(12), 2020

work page 2020

[27] [27]

Relative rates of surface and volume synthesis set bacterial cell size

Leigh K Harris and Julie A Theriot. Relative rates of surface and volume synthesis set bacterial cell size. Cell, 165(6):1479–1492, 2016

work page 2016

[28] [28]

The fission yeast cell size control system integrates pathways measuring cell surface area, volume, and time

Kristi E Miller, Cesar Vargas-Garcia, Abhyudai Singh, and James B Moseley. The fission yeast cell size control system integrates pathways measuring cell surface area, volume, and time. Current Biology , 33(16):3312–3324, 2023

work page 2023

[29] [29]

Size-dependent expres- sion of the fission yeast cdc13 cyclin is conferred by trans- lational regulation

Samirul Bashir, Xi-Ming Sun, Yixuan Zhao, Nuria G Mart´ ınez-Illescas, Isabella Gallego-L´ opez, Lauren Guer- rero Negr´ on, Daniel Keifenheim, Tatiana Karadimitriou, Thi Tran, Mary Pickering, et al. Size-dependent expres- sion of the fission yeast cdc13 cyclin is conferred by trans- lational regulation. bioRxiv, pages 2023–01, 2023

work page 2023

[30] [30]

Pom1 and cell size homeostasis in fission yeast.Cell cycle, 12(19):3417–3425, 2013

Elizabeth Wood and Paul Nurse. Pom1 and cell size homeostasis in fission yeast.Cell cycle, 12(19):3417–3425, 2013

work page 2013

[31] [31]

See Supplemental Material for additional derivations, and stochastic simulations

work page

[32] [32]

Dilution of the cell cycle inhibitor whi5 con- trols budding-yeast cell size

Kurt M Schmoller, JJ Turner, M K˜ oivom¨ agi, and Jan M Skotheim. Dilution of the cell cycle inhibitor whi5 con- trols budding-yeast cell size. Nature, 526(7572):268–272, 2015

work page 2015

[33] [33]

Control of cell cycle transcription during g1 and s phases

Cosetta Bertoli, Jan M Skotheim, and Robertus AM De Bruin. Control of cell cycle transcription during g1 and s phases. Nature reviews Molecular cell biology , 14(8):518–528, 2013

work page 2013

[34] [34]

Rb and cell cycle progression

C Giacinti and Antonion Giordano. Rb and cell cycle progression. Oncogene, 25(38):5220–5227, 2006

work page 2006

[35] [35]

Protein turnover forms one of the highest maintenance costs in lactococcus lactis

Petri-Jaan Lahtvee, Andrus Seiman, Liisa Arike, Kaarel Adamberg, and Raivo Vilu. Protein turnover forms one of the highest maintenance costs in lactococcus lactis. Microbiology, 160(7):1501–1512, 2014

work page 2014

[36] [36]

Evolution- ary advantage of cell size control

Spencer Hobson-Gutierrez and Edo Kussell. Evolution- ary advantage of cell size control. Physical Review Let- ters, 134(11):118401, 2025

work page 2025

[37] [37]

From noisy cell size control to popula- tion growth: When variability can be beneficial

Arthur Genthon. From noisy cell size control to popula- tion growth: When variability can be beneficial. Physical Review E, 111(3):034407, 2025. 1 SUPPLEMENT AL MA TERIAL CONCENTRA TION DYNAMICS Here, we solve Eq. 3 for both exponential and linear size growth. We focus first on the exponential size growth case (s = bneαTn , ρ = 1/αbn) where ∂c(bn, Tn) ∂bn...

work page 2025