Pith. sign in

REVIEW 3 major objections 4 minor 78 references

Growth and remodeling control shape memory in morphogenetic rods

T0 review · 3 major / 4 minor · reviewed 2026-07-31 · deepseek-v4-flash

Pith's one-line read Growth breaks the trade-off between shape complexity and reproducibility in remodeling rods.

desk verdict A clean model-level phase diagram for shape memory in growing visco-elasto-plastic rods, with one genuine stochastic-projection gap in the high-plasticity branch that likely shifts boundaries but not the qualitative story. read the letter →

arxiv 2607.23907 v1 pith:CF4GY7PP submitted 2026-07-27 cond-mat.soft physics.bio-phq-bio.TO

classification cond-mat.softphysics.bio-phq-bio.TO
keywords shapememorymorphogenesiselasticaplasticitygrowthbucklingmodesnoiseviscoelasticrods
verification ladder T0 review T1 audit T2 compute T3 formal

The pith

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

The reading

Elastic slender rods buckle into the simplest possible shape, so a complex initial pattern tends to coarsen away. This paper shows that internal remodeling (plasticity) can freeze that coarsening, but at high rates plasticity makes the rod's rest shape follow the current shape so closely that noise drives the pattern to diffuse away. The trade-off between these two failure modes leaves only a window of intermediate plasticity for shape memory. Uniform growth changes both: it slows elastic coarsening and dilutes noise, so with fast growth and high plasticity, complex initial patterns remain reproducible. The results frame remodeling rate and growth rate as two control knobs for whether an encoded shape is remembered, degraded, or transformed.

What carries the argument

The central object is the plastica model: a one-dimensional elastica with internal viscosity µ, rest-curvature remodeling rate η, and an end-shortening constraint that makes the tension a Lagrange multiplier. In mode space the dynamics reduce to a replicator-like competition among bending modes (with decaying selection strength e^{−ηt}) at low plasticity, and to anisotropic Brownian motion on the constraint sphere at high plasticity, with mode-dependent mobility γ_n = µ + Bq_n²/η. Two dimensionless numbers carry the argument: the plasticity number Pl = ητ_E and the normalized growth rate gτ_E. The work also supplies explicit formulas for the coarsening threshold, the diffusion memory time t_

What would settle it

Simulate the full plastica at large plasticity without dropping the noise term in the tension equation; if the pattern is lost at a different time or growth rate than predicted by the F=0 formula, the trade-off-breaking claim fails. Alternatively, a physical experiment on a growing, remodeling gel rod that shows coarsening at high growth rather than preservation would refute it.

Watch

Extended reading notes

Core claim

The paper establishes that in a minimal visco-elasto-plastic rod—the 'plastica'—the fate of an initially encoded multi-mode shape depends on two dimensionless numbers: the plasticity number Pl = ητ_E and the normalized growth rate gτ_E. At low plasticity, bending modes compete like replicators with fitness proportional to −q_n², so the lowest mode excludes all others and erases complexity. Moderately fast remodeling arrests that coarsening because the rest curvature tracks the current shape before selection completes, protecting the pattern. At high plasticity, the rod behaves as a nearly fluid thread: the rest shape adiabatically follows the current shape, the constraint force vanishes at l

Load-bearing premise

The predictions for how noise destroys a pattern at high plasticity rest on assuming the rod's tension is effectively zero once the rest shape tracks the current shape; if that tension is not negligible, the boundary between remembered and lost patterns changes.

Editorial extensions

If this is right

  • Below a minimum plasticity, any multi-mode pattern deterministically coarsens to the fundamental buckling mode, so complexity cannot be encoded without remodeling.
  • Above a second, higher plasticity, noise converts into geometric disorder and the pattern is lost after a plasticity-dependent memory time; the window of successful memory shrinks for more complex initial shapes.
  • Growth shifts both boundaries: even at zero plasticity, sufficiently fast growth suppresses coarsening, and sufficiently fast growth makes noise diffusion unable to erase the pattern.
  • In the high-growth, high-plasticity quadrant, arbitrary initial conditions (including random ones) produce reproducible, complex shapes, providing a design rule for morphogenesis and for fluctuation-tolerant soft structures.
  • The theory makes quantitative predictions for biological systems: an early constriction in a developing gut should be amplified into a stereotyped loop only within a finite window of remodeling rates, a window that growth widens.

Reading between the lines

Editorial extensions of the paper, not claims the author makes directly.

  • The replicator-dynamics mapping suggests the same trade-off may appear in any adaptive system where competing components have frequency-dependent fitness and a decaying selection strength—for example, regulatory networks with plasticity—so the 'growth suppresses both failure modes' result might generalize beyond rods.
  • The paper treats internal viscosity only; the supplement's external-viscosity version shows intermediate coarsening transients, hinting that the phase diagram depends on where dissipation acts—an extension worth exploring for tissues immersed in fluid.
  • A direct experimental test could use a swelling or growing gel rod with tunable remodeling rate and measure the Pearson correlation of mode spectra over time; the predicted non-monotonic memory with plasticity, and the rescuing effect of growth, are observable.
  • The assumption of constant bending rigidity B during growth is conservative; if the rod thins as it elongates, coarsening would be suppressed even faster, so the qualitative result is robust but the quantitative thresholds would shift.
Share X Bluesky LinkedIn Reddit HN

Signed reviews

No signed human review yet.

Editorial analysis

A structured set of objections, weighed in public.

Desk editor's note, referee report, and a circularity audit.

Referee Report

3 major / 4 minor

Summary. The paper asks when an initially patterned shape is remembered during growth and remodeling in a slender rod. The authors study a minimal visco-elasto-plastic 'plastica' model in the small-slope limit: tangent-angle modes evolve under overdamped elasticity and a rest-curvature field that relaxes to the current shape at rate η. Two failure modes are identified. At low plasticity the deterministic dynamics are replicator-like and coarse-grain to the fundamental mode; above a plasticity threshold, initial high-mode patterns are protected. At high plasticity the rod becomes fluid-like and noise drives anisotropic diffusion on the constraint sphere, erasing the pattern over a memory time t_D. Growth, modeled as uniform elongation, reduces both the elastic coarsening rate and the effective noise amplitude, and the authors derive criteria for a growth rate above which complex patterns are retained despite noise. The central claim is that growth breaks the plasticity trade-off between complexity and reproducibility. The paper includes analytical derivations in the Supplement, small-angle spectral simulations, fully-nonlinear checks, and a code repository.

Significance. If the quantitative phase boundaries are correct, this is a valuable conceptual result: it identifies two dimensionless knobs, plasticity and growth rate, that control whether an encoded mechanical pattern is remembered, degraded, or transformed. The model is deliberately minimal, and the deterministic coarsening derivation (Supplement II.A) is clean. The thresholds have no fitted parameters: ε comes from the initial perturbation and ε_σ from the noise strength, so the agreement with simulations is a genuine internal consistency check. The paper goes beyond linear stability by checking against fully-nonlinear finite-difference simulations and by considering an external-viscosity variant with similar phenomenology. These strengths make the manuscript a good candidate for a broadly read soft-matter journal, provided the high-plasticity stochastic projection is made rigorous or numerically verified.

major comments (3)
  1. [Supplement II.B, Eq. (S33)] The high-plasticity boundary, t_D, the criterion Φ(T_g)≤1, and g∞ all rely on the statement that the constraint force vanishes after differentiating Σ_n θ_n^2=C while 'ignoring the noise'. But from γ_n θdot_n = F θ_n + √(2σ/L) ζ_n, exact constraint enforcement gives F = −√(2σ/L) (Σ_n θ_n ζ_n/γ_n)/(Σ_n θ_n^2/γ_n), not F=0. Dropping the noise in this step is not controlled: white noise is unbounded, and after the θ_nF term is expanded the noise-induced drift is of order σ, the same order as the diffusion term. The authors' own remark in Supplement II.C that the tension does not vanish exactly at lower Pl with noise is direct evidence that the correction is not identically zero. Request either a proper stochastic projection (which may introduce an O(σ) drift that renormalizes d_eff and t_D) or direct numerical measurement of t_D and g∞ at the Pl values where the boundary is drawn, to confir
  2. [Main text, Eq. (5) and Fig. 2c] The coarsening-protection threshold in the presence of noise is obtained by replacing the deterministic initial perturbation amplitude ε in Eq. (5) with ε_σ = σ/(μBq_1^2 C). This replacement is plausible but is not derived from the stochastic dynamics; it treats noise as if it were a static initial condition. Since the lower boundary of the 'preserved' region is one of the paper's main quantitative predictions, this step needs support, either by a stochastic multiscale derivation or by showing that the boundary coincides with a measured escape/fixation time from the basin of the initial mode. As written, the lower boundary is an ansatz rather than a consequence of the equations.
  3. [Supplement III.B.2, Eq. (S50)] The growth-modified diffusion integral replaces d_eff(t) by d_eff(0). Since L(t) increases, q_n(t)=πn/L(t) decreases and d_eff(t) increases with time, so this approximation is not conservative at finite Pl: it underestimates the total angular drift Φ(T_g) and can therefore overestimate the critical growth rate g∞. At Pl→∞ the error vanishes because d_eff→d, and the authors state that the large-η limit is the regime of interest, but the text should state the approximation explicitly and give a bound or numerical check for finite Pl. This is a local fix, not a challenge to the central claim.
minor comments (4)
  1. [Notation, Eq. (1) and Fig. 1] The elastic timescale is defined as τ_E=μL_0^2/B, but some passages write the elastic rate as B/(μL^2) with L appearing to be the current length. Please use a consistent symbol (L_0, L(t), or L_1) in the definitions of Pl and in the thresholds.
  2. [Fig. 2d] The colorbar is cropped at m=12 with a '>12' label, while the text says the highest mode can go up to d=64. This makes the m-dependence of the preserved window difficult to read; consider plotting the full range or stating the crop explicitly in the caption.
  3. [Supplement III.B.2, Eq. (S50)] The replacement d_eff(t)≈d_eff(0) is introduced with an unqualified '≈'. Since this approximation enters a published design rule, a sentence should state the size of the correction and why it is negligible in the regime shown in Fig. 3.
  4. [References] Reference [42] is listed as 'To be submitted'. If it is not yet available, it should be marked as unpublished work or removed from the numbered reference list; if it is a preprint, a stable identifier is needed.

Circularity Check

0 steps flagged · score 0.0 of 10

No significant circularity: the analytic phase boundaries are derived from the model equations with no fitted parameters and checked against separate simulations.

full rationale

The derivation chain is self-contained. The low-plasticity coarsening threshold follows from integrating the mode-fraction amplification (Eq. 4) to obtain the plasticity criterion (Eq. 5); the high-plasticity diffusion boundary follows from the large-η effective dynamics (Supp. Eq. S32) and the spherical constraint, giving t_D and g∞ without fitting parameters; the growth criteria follow from the same integrals with L(t) (Supp. Eqs. S38, S41, S50-S52). The only parameter replacing the perturbation in the noisy case, ε_σ, is computed from σ and the model parameters, not fit to simulation outcomes. The main caveat—Supp. II.B obtains F=0 by differentiating the constraint while 'ignoring the noise' (Eq. S33)—is an approximation, and the Supplement itself notes the spectra do not perfectly collapse 'due to the tension F not vanishing exactly at lower Pl in the presence of noise' (Supp. II.C). That is a correctness/robustness risk, not a circular reduction: the 'lost' region and growth-rescue threshold are not defined by the simulation measurements or by a fitted parameter. Self-citations [35,41,42] are contextual biological examples rather than load-bearing support, and [49] is the paper's own Supplement. Thus no prediction reduces by construction to its input. Score 0.

Assumptions & free parameters 0 free parameters · 8 assumptions · 0 invented entities

No parameter is fitted to data. Pl, g*tau_E, sigma_bar, Delta/L0, L1/L0, Lf/L1, d, and m are physical or numerical control parameters whose values are stated and varied; the theoretical boundaries are functions of these inputs, not adjustable fits. No new physical entities are postulated; 'plastica' is the name of the model, not a new entity.

assumptions (8)
  • domain assumption Constitutive law M=B(theta'-phi') and linear rest-curvature relaxation dphi'/dt=eta(theta'-phi') (Eq. 1) capture the mechanics of morphodynamic rods.
    The entire paper is built on this visco-elasto-plastic rod model; no stress-dependent or nonlinear remodeling is considered.
  • domain assumption Overdamped dynamics with internal viscosity mu dominate inertia and external drag (Eq. 1a).
    The overdamped Langevin equation is asserted in the main text; the external-viscosity variant is treated only in Supplement IV.
  • domain assumption Small-angle approximation |theta|<<1 and Delta/L0<<1; Eq. (1) is the small-Delta limit of the fully nonlinear formulation.
    The main text restricts to Delta<0.1 L0 and states agreement with the fully nonlinear formulation; the simulation parameters sit near the edge of this regime.
  • domain assumption Noise is additive white Gaussian with constant amplitude sigma, independent of mode, plasticity, and growth.
    Used in Eq. (1a) and mode-space Eq. (2a); the high-plasticity diffusion theory uses this to compute d_eff and t_D.
  • domain assumption Finite Fourier truncation to d modes, with d~L0/h set by the rod aspect ratio.
    d determines d_eff and the growth threshold g_infinity; the optimal plasticity window shrinks as d grows.
  • domain assumption Uniform exponential growth L(t)=L1 exp(g t) with constant bending modulus B.
    Used to derive g0 and g_infinity in Supplement III; the paper notes other growth laws or thinning would change quantitative thresholds.
  • standard math Watson's lemma expansion phi_n=theta_n-eta^{-1} dtheta_n/dt + O(eta^{-2}) is valid in the large-eta regime.
    Supplement II.B, Eqs. (S30)-(S31); standard asymptotic expansion, and the resulting F=0 is tested against numerics.
  • standard math Replicator-equation form of mode competition in the elastic limit is exact under the small-angle constraint.
    Supplement II.A, Eq. (S28); uses the known replicator equation from population dynamics (Nowak 2006; Schuster and Sigmund 1983).

how reviews work

0 comments
Cite this review

Pith. "Pith review of Growth and remodeling control shape memory in morphogenetic rods." pith.science (2026). https://pith.science/paper/CF4GY7PP

@misc{pith2026260723907,
  author       = {Pith},
  title        = {Pith review of: Growth and remodeling control shape memory in morphogenetic rods},
  year         = {2026},
  howpublished = {\url{https://pith.science/paper/CF4GY7PP}},
  note         = {Machine review of arXiv:2607.23907}
}
read the original abstract

Mechanical instabilities provide a general design principle for shaping developing organs and engineering soft materials. However, in slender structures, simple elastic buckling tends to erase rather than preserve shape complexity: structures relax to the simplest possible shape, erasing finer detail. Living systems nonetheless build complex, reproducible morphologies from continually remodeling material, while remaining robust to noise arising across scales. Using analytical theory and numerical simulations of a minimal model of growing visco-elasto-plastic rods, we show that remodeling plays two opposing roles: At low plasticity, patterns coarsen through elastic relaxation, while high plasticity converts fluctuations into geometric disorder. This sets a trade-off between shape complexity and reproducibility with an optimal intermediate plasticity, which protects initial patterns. Growth breaks this trade-off by suppressing both failure modes, enabling complex shapes to be reproducibly generated. Our results identify remodeling and growth rates as two knobs governing whether an encoded pattern is remembered, degraded, or transformed.

Figures

Figures reproduced from arXiv: 2607.23907 by the authors.

Figure 1
Figure 1. FIG. 1. Does complex rheology stabilize noisy morphogene [PITH_FULL_IMAGE:figures/full_fig_p001_1.png] view at source ↗
Figure 2
Figure 2. FIG. 2. Plasticity suppresses deterministic coarsening but induces noise-driven state diffusion. (a) In the absence of growth, [PITH_FULL_IMAGE:figures/full_fig_p003_2.png] view at source ↗
Figure 3
Figure 3. FIG. 3. Growth stabilizes patterns. With an initial condition [PITH_FULL_IMAGE:figures/full_fig_p004_3.png] view at source ↗

Discussion (0). Continue with ORCID to comment.

Reference graph

Works this paper leans on

78 extracted references · 1 canonical work pages

  1. [1]

    Savin, N

    T. Savin, N. A. Kurpios, A. E. Shyer, P. Florescu, H. Liang, L. Mahadevan, and C. J. Tabin, On the growth and form of the gut, Nature476, 57 (2011)

  2. [2]

    A. E. Shyer, T. Tallinen, N. L. Nerurkar, Z. Wei, E. S. Gil, D. L. Kaplan, C. J. Tabin, and L. Mahadevan, Vil- lification: How the Gut Gets Its Villi, Science342, 212 (2013)

  3. [3]

    Capolupo, C

    L. Capolupo, C. Baader, P. Robin, S. Barbiero, K. C. Oost, J. Timmer, S. Suppinger, Q. Yang, V. Kalck, A.-M. Lennon-Dumenil, E. Hannezo, and P. Liberali, Tis- sue swelling and local mechanosensing drive crypt con- vergence in intestinal development (2026), hal preprint

  4. [4]

    S. J. Gerbode, J. R. Puzey, A. G. McCormick, and L. Ma- hadevan, How the Cucumber Tendril Coils and Over- winds, Science337, 1087 (2012)

  5. [5]

    D. E. Moulton, H. Oliveri, and A. Goriely, Multiscale in- tegration of environmental stimuli in plant tropism pro- duces complex behaviors, Proceedings of the National Academy of Sciences117, 32226 (2020)

  6. [6]

    Armon, E

    S. Armon, E. Efrati, R. Kupferman, and E. Sharon, Ge- ometry and Mechanics in the Opening of Chiral Seed Pods, Science333, 1726 (2011)

  7. [7]

    Si´ efert, E

    E. Si´ efert, E. Reyssat, J. Bico, and B. Roman, Program- ming curvilinear paths of flat inflatables, Proceedings of the National Academy of Sciences116, 16692 (2019)

  8. [8]

    Pezzulla, S

    M. Pezzulla, S. A. Shillig, P. Nardinocchi, and D. P. Holmes, Morphing of geometric composites via residual swelling, Soft Matter11, 5812 (2015)

Show all 78 references
  1. [9]

    Y. Sun, W. M. Choi, H. Jiang, Y. Y. Huang, and J. A. Rogers, Controlled buckling of semiconductor nanorib- bons for stretchable electronics, Nature Nanotechnology 1, 201 (2006)

  2. [10]

    Audoly and Y

    B. Audoly and Y. Pomeau,Elasticity and geometry: from hair curls to the non-linear response of shells, first pub- lished in paperback ed. (Oxford University Press, Oxford, 2018)

  3. [11]

    S. S. Antman,Nonlinear Problems of Elasticity, Applied Mathematical Sciences, Vol. 107 (Springer-Verlag, New York, 2005)

  4. [12]

    Sydney Gladman, E

    A. Sydney Gladman, E. A. Matsumoto, R. G. Nuzzo, L. Mahadevan, and J. A. Lewis, Biomimetic 4D printing, Nature Materials15, 413 (2016)

  5. [13]

    W. M. van Rees, E. Vouga, and L. Mahadevan, Growth patterns for shape-shifting elastic bilayers, Proceedings of the National Academy of Sciences114, 11597 (2017)

  6. [14]

    J. W. Boley, W. M. van Rees, C. Lissandrello, M. N. Horenstein, R. L. Truby, A. Kotikian, J. A. Lewis, and L. Mahadevan, Shape-shifting structured lattices via multimaterial 4D printing, Proceedings of the National Academy of Sciences116, 20856 (2019)

  7. [15]

    Klein, E

    Y. Klein, E. Efrati, and E. Sharon, Shaping of Elastic Sheets by Prescription of Non-Euclidean Metrics, Science 315, 1116 (2007)

  8. [16]

    Zhang, M

    Y. Zhang, M. Moshe, and E. Sharon, Isometric incompat- ibility in growing elastic sheets (2026), arXiv:2603.21112 [cond-mat.soft]

  9. [17]

    N. Gov, A. G. Zilman, and S. Safran, Cytoskeleton Con- finement and Tension of Red Blood Cell Membranes, Physical Review Letters90, 228101 (2003)

  10. [18]

    Koˇ smrlj and D

    A. Koˇ smrlj and D. R. Nelson, Statistical Mechanics of Thin Spherical Shells, Physical Review X7, 011002 (2017)

  11. [19]

    Radja, E

    A. Radja, E. M. Horsley, M. O. Lavrentovich, and A. M. Sweeney, Pollen Cell Wall Patterns Form from Modu- lated Phases, Cell176, 856 (2019)

  12. [20]

    J. A. Jackson, N. Romeo, A. Mietke, K. J. Burns, J. F. Totz, A. C. Martin, J. Dunkel, and J. Imran Alsous, Scal- ing behaviour and control of nuclear wrinkling, Nature Physics19, 1927 (2023)

  13. [21]

    Ambrosi, M

    D. Ambrosi, M. Ben Amar, C. J. Cyron, A. DeSimone, A. Goriely, J. D. Humphrey, and E. Kuhl, Growth and remodelling of living tissues: perspectives, challenges and opportunities, Journal of The Royal Society Interface16, 20190233 (2019)

  14. [22]

    Goriely,The Mathematics and Mechanics of Biological Growth, Interdisciplinary Applied Mathematics, Vol

    A. Goriely,The Mathematics and Mechanics of Biological Growth, Interdisciplinary Applied Mathematics, Vol. 45 (Springer New York, New York, NY, 2017)

  15. [23]

    Moulton, T

    D. Moulton, T. Lessinnes, and A. Goriely, Morphoelastic rods. Part I: A single growing elastic rod, Journal of the Mechanics and Physics of Solids61, 398 (2013)

  16. [24]

    D. S. Alber, S. Zhao, A. O. Jacinto, E. F. Wieschaus, S. Y. Shvartsman, and P. A. Haas, A model for boundary- driven tissue morphogenesis, Proceedings of the National Academy of Sciences122, e2505160122 (2025)

  17. [25]

    Chopin, M

    J. Chopin, M. Dasgupta, and A. Kudrolli, Dynamic Wrin- kling and Strengthening of an Elastic Filament in a Vis- cous Fluid, Physical Review Letters119, 088001 (2017)

  18. [26]

    Kodio, A

    O. Kodio, A. Goriely, and D. Vella, Dynamic buckling of an inextensible elastic ring: Linear and nonlinear analy- ses, Physical Review E101, 053002 (2020)

  19. [27]

    J. L. Shivers, M. Nguyen, A. R. Dinner, P. M. Vla- hovska, and S. Vaikuntanathan, Renormalized Mechan- ics and Stochastic Thermodynamics of Growing Vesicles, 6 PRX Life4, 013012 (2026)

  20. [28]

    D. A. Matoz-Fernandez, F. A. Davidson, N. R. Stanley- Wall, and R. Sknepnek, Wrinkle patterns in active vis- coelastic thin sheets, Physical Review Research2, 013165 (2020)

  21. [29]

    S. C. Al-Izzi, G. Rowlands, P. Sens, and M. S. Turner, Hydro-osmotic Instabilities in Active Membrane Tubes, Physical Review Letters120, 138102 (2018)

  22. [30]

    Y. Liu, B. Chakrabarti, D. Saintillan, A. Lindner, and O. Du Roure, Morphological transitions of elas- tic filaments in shear flow, Proceedings of the National Academy of Sciences115, 9438 (2018)

  23. [31]

    A. P. Singh and C. N¨ usslein-Volhard, Zebrafish Stripes as a Model for Vertebrate Colour Pattern Formation, Cur- rent Biology25, R81 (2015)

  24. [32]

    Corson, L

    F. Corson, L. Couturier, H. Rouault, K. Mazouni, and F. Schweisguth, Self-organized Notch dynamics generate stereotyped sensory organ patterns inDrosophila, Science 356, eaai7407 (2017)

  25. [33]

    Slavkov, D

    I. Slavkov, D. Carrillo-Zapata, N. Carranza, X. Diego, F. Jansson, J. Kaandorp, S. Hauert, and J. Sharpe, Morphogenesis in robot swarms, Science Robotics3, eaau9178 (2018)

  26. [34]

    L. S. Tsimring, Noise in biology, Reports on Progress in Physics77, 026601 (2014)

  27. [35]

    Romeo, D

    N. Romeo, D. G. Martin, M. Scandolo, M. Fruchart, E. M. Munro, and V. Vitelli, Information bounds the robustness of self-organized systems (2026), arXiv:2511.01682

  28. [36]

    Mongera, P

    A. Mongera, P. Rowghanian, H. J. Gustafson, E. Shelton, D. A. Kealhofer, E. K. Carn, F. Serwane, A. A. Lucio, J. Giammona, and O. Camp` as, A fluid-to-solid jamming transition underlies vertebrate body axis elongation, Na- ture561, 401 (2018)

  29. [37]

    N. I. Petridou, B. Corominas-Murtra, C.-P. Heisenberg, and E. Hannezo, Rigidity percolation uncovers a struc- tural basis for embryonic tissue phase transitions, Cell 184, 1914 (2021)

  30. [38]

    Raspopovic, L

    J. Raspopovic, L. Marcon, L. Russo, and J. Sharpe, Digit patterning is controlled by a Bmp-Sox9-Wnt Turing net- work modulated by morphogen gradients, Science345, 566 (2014)

  31. [39]

    Serafini, M

    G. Serafini, M. Setoudeh, M. B. Cuenca, C. Brillard, M. Arzt, P. Mejstˇ rik, P. A. Haas, and P. Tomanˇ c´ ak, Embryo-eggshell interaction counteracts chiral bias in early Drosophila morphogenesis (2026)

  32. [40]

    C. M. Smits, S. Dutta, V. Jain-Sharma, S. J. Streichan, and S. Y. Shvartsman, Maintaining symmetry during body axis elongation, Current Biology33, 3536 (2023)

  33. [41]

    N. P. Mitchell, D. J. Cislo, S. Shankar, Y. Lin, B. I. Shraiman, and S. J. Streichan, Visceral organ morpho- genesis via calcium-patterned muscle constrictions, eLife 11, e77355 (2022)

  34. [42]

    Romeo, C

    N. Romeo, C. Anto, A. Strok, E. L. Hendricks, N. A. Tamarina, F. Brauns, and N. P. Mitchell, To be submit- ted

  35. [43]

    R. E. Goldstein and A. Goriely, Dynamic buckling of morphoelastic filaments, Physical Review E74, 010901 (2006)

  36. [44]

    Howell, G

    P. Howell, G. Kozyreff, and J. Ockendon,Applied Solid Mechanics, 1st ed. (Cambridge University Press, 2001)

  37. [45]

    Bergou, B

    M. Bergou, B. Audoly, E. Vouga, M. Wardetzky, and E. Grinspun, Discrete viscous threads, ACM Transac- tions on Graphics29, 1 (2010)

  38. [46]

    Audoly, N

    B. Audoly, N. Clauvelin, P.-T. Brun, M. Bergou, E. Grin- spun, and M. Wardetzky, A discrete geometric approach for simulating the dynamics of thin viscous threads, Jour- nal of Computational Physics253, 18 (2013)

  39. [47]

    P.-T. Brun, B. Audoly, N. M. Ribe, T. S. Eaves, and J. R. Lister, Liquid Ropes: A Geometrical Model for Thin Viscous Jet Instabilities, Physical Review Letters114, 174501 (2015)

  40. [48]

    Kamrin and L

    K. Kamrin and L. Mahadevan, Soft catenaries, Journal of Fluid Mechanics691, 165 (2012)

  41. [49]

    See Supplemental Material below for derivations and ad- ditional simulations

  42. [50]

    A. C. Martin, M. Kaschube, and E. F. Wieschaus, Pulsed contractions of an actin–myosin network drive apical con- striction, Nature457, 495 (2009)

  43. [51]

    M. A. Nowak,Evolutionary dynamics: exploring the equations of life(The Belknap Press of Harvard Univer- sity Press, Cambridge, Massachusetts London, England, 2006)

  44. [52]

    The correlationρis taken using the covariance over all modes

  45. [53]

    J. R. Gladden, N. Z. Handzy, A. Belmonte, and E. Viller- maux, Dynamic Buckling and Fragmentation in Brittle Rods, Physical Review Letters94, 035503 (2005)

  46. [54]

    C. M. Nelson, On Buckling Morphogenesis, Journal of Biomechanical Engineering138, 021005 (2016)

  47. [55]

    D. E. Ingber, Mechanical control of tissue morphogene- sis during embryological development, The International Journal of Developmental Biology50, 255 (2006)

  48. [56]

    N. I. Petridou and C. Heisenberg, Tissue rheology in embryonic organization, The EMBO Journal38, EMBJ2019102497 (2019)

  49. [57]

    Tallinen, J

    T. Tallinen, J. Y. Chung, F. Rousseau, N. Girard, J. Lef` evre, and L. Mahadevan, On the growth and form of cortical convolutions, Nature Physics12, 588 (2016)

  50. [58]

    G. P. Choi, C. Liu, S. Yin, G. S´ ejourn´ e, R. S. Smith, C. A. Walsh, and L. Mahadevan, Biophysical basis for brain folding and misfolding patterns in ferrets and humans, eLife14, RP107141 (2025)

  51. [59]

    L. A. Hoffmann and L. Mahadevan, How to grow a straight filament (2026), arXiv:2606.11080 [cond- mat.soft]

  52. [60]

    Imran Alsous, B

    J. Imran Alsous, B. Chakrabarti, B. Palmer, and M. J. Shelley, The physical consequences of sperm gigantism, Nature Physics 10.1038/s41567-026-03305-4 (2026)

  53. [61]

    T˘ atulea-Codrean and E

    M. T˘ atulea-Codrean and E. Lauga, Elastohydrodynamic Synchronization of Rotating Bacterial Flagella, Physical Review Letters128, 208101 (2022)

  54. [62]

    Schuster and K

    P. Schuster and K. Sigmund, Replicator dynamics, Jour- nal of Theoretical Biology100, 533 (1983)

  55. [63]

    Cressman and Y

    R. Cressman and Y. Tao, The replicator equation and other game dynamics, Proceedings of the National Academy of Sciences111, 10810 (2014)

  56. [64]

    Stern, M

    M. Stern, M. Guzman, F. Martins, A. J. Liu, and V. Bal- asubramanian, Physical Networks Become What They Learn, Physical Review Letters134, 147402 (2025)

  57. [65]

    Supplement to: Growth and remodeling control shape memory in morphogenetic rods Nicolas Romeo, 1, 2, 3, 4,∗ David B

    Code is available athttps://github.com/ MitchellLabCode/plastica. Supplement to: Growth and remodeling control shape memory in morphogenetic rods Nicolas Romeo, 1, 2, 3, 4,∗ David B. Br¨ uckner,5, 6 and Noah P. Mitchell 3, 7, 1, 4,† 1Center for Living Systems, University of Ch...

  58. [66]

    Small-angle formulation 3

  59. [67]

    Pearson correlation 5 II

    Fully nonlinear formulation 3 D. Pearson correlation 5 II. Derivation of memory criteria in the absence of growth 5 A. Derivation of replicator equations 5 B. Derivation of large plasticity effective dynamics 5 C. Steady-state spectrum 7 III. Derivation of memory criterion for...

  60. [68]

    Growth in the large-plasticity regime produces limited coarsening 9

  61. [69]

    Alternative model with external viscosity 10 References 11 I

    State diffusion with growth 9 IV. Alternative model with external viscosity 10 References 11 I. PLASTICA A. F ully-nonlinear formulation We consider the elastica system, with a constitutive equationM(s, t) =B(θ ′(s, t)−ϕ′(s, t)), whereθis defined by the tangent vector to the c...

  62. [70]

    Small-angle formulation We use a spectral semi-implicit scheme to solve the linearized scheme: the tension is computed explicitly, but the timestepping of the ODEs is done implicitly. More precisely, at timestepkwe computeT k =µ ˙C(t k)/C(t k) + B C(tk) P n θn(tk)(θn(tk)−ϕ n(t...

  63. [71]

    The arclength domains∈[0, L(t)] is sampled atN pts equispaced points si = (i−1)δ, i= 1,

    Fully nonlinear formulation The fully nonlinear dynamics is solved using a finite-difference approximation of the differential operators, with a nonlinear implicit Euler time stepper which requires iterative Newton solves. The arclength domains∈[0, L(t)] is sampled atN pts equ...

  64. [72]

    UpdateL n+1 =L 0eg tn+1 ,δ n+1 =L n+1/(Npts −1), and the Simpson weightsw int i from (S19)

  65. [73]

    Add noiseθ n i ←θ n i +σ iN(0,1) via (S22), with the newδ n+1 inV i

  66. [74]

    Newton iterate (S21) until the residual|R| ∞ <tolerance (∼10 −8)

  67. [75]

    species” are the bending modes, andr n =θ 2 n/C is the fraction of the “population

    Recoverϕ n+1 from (S20). In practice, we find that this solver is consistent with the small-angle spectral solver for perturbations of amplitude ϵ >10 −6 in deterministic simulations (Fig. S1). At smallerϵ, the limited precision of this solver which estimates spatial derivativ...

  68. [76]

    Growth in the large-plasticity regime produces limited coarsening The growth-induced nonzero tension leads the deterministic dynamics to obey a replicator equation with fitness 1/γn, ˙rn = 2 C θn ˙θn −θ 2 n ˙C C 2 (S45) = ˙C C 1/γn ⟨γ−1⟩ −1 rn (S46) driving the mode fraction t...

  69. [77]

    We now compute the timescale of diffusion with g >0

    State diffusion with growth The results above indicate that we can still ignore the effect of deterministic coarsening at large plasticity, which is then still dominated by the effect of stochastic mode diffusion. We now compute the timescale of diffusion with g >0. In the pre...

  70. [78]

    Edwards.The theory of polymer dynamics

    Masao Doi and Samuel F. Edwards.The theory of polymer dynamics. Number 73 in International series of monographs on physics. Clarendon Press, Oxford, reprint edition, 2013

Pith tools

Reviewed July 31, 2026 · model on record in the stance chip above.