Mechanical constraints influence microtubule orientation in plant cells
Bibliographic record
Abstract
ABSTRACT In growing plant cells, parallel ordering of microtubules (MTs) along the inner surface of the cell membrane influences the direction of cell expansion and thereby plant morphology. For correct expansion of organs that primarily grow by elongating, such as roots and stems, MTs must bend in the high-curvature direction along the cylindrically shaped cell membrane in order to form the required circumferential arrays. Previous studies, which have recapitulated the self-organization of these arrays, ignored MT mechanics and assumed MTs follow geodesics of the cell surface. Here, we show, through analysis of a derived Euler-Lagrange equation, that an elastic MT constrained to a cylindrical surface will deflect away from geodesics and toward low curvature directions to minimize bending energy. This occurs when the curvature of the cell surface is relatively high for a given anchor density. In the limit of infinite anchor density, MTs always follow geodesics. We compare our analytical predictions to measured curvatures and anchor densities and find that the regime in which cells are forming these cortical arrays straddles the region of parameter space in which arrays must form under the antagonistic influence of this mechanically induced deflection. Although this introduces a potential obstacle to forming circumferentially orientated arrays that needs to be accounted for in the models, it also raises the question of whether plants use this mechanical phenomenon to regulate array orientation. The model also constitutes an elegant generalization of the classical Euler-bucking instability along with an intrinsic unfolding of the associated pitchfork bifurcation. SIGNIFICANCE STATEMENT The cortical microtubule array in plant cells is a dynamic structure whose organization influences cell function, growth, and plant morphology. During cell growth, the array must be highly ordered and perpendicular to the axis of expansion. In other instances, it can be disordered or axially oriented. Experimental and modelling studies have outlined the importance of various microtubule-microtubule interactions in the ordering and orienting process. We explore the influence of cell shape on the bending of microtubules that has been largely ignored to date. We find that cell shape can induce microtubule bending that steers them away from high-curvature directions. This bending force can act as an obstacle to proposed mechanisms for array organization and also as a mechanism for regulating reorientation of the array.
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How this classification was reachedexpand
Full frame machine prediction
Teacher imitationNot calibrated prevalence, not ground truth. Human validation pending. The Gemma side is a direct model label for every work in the frame, read from the title-only record. The Codex side is a classifier learned from the 10,348 direct Codex labels and calibrated to design-weighted sample rates; fields without enough sample support carry no Codex call. Candidate is the union of the two sides; consensus is their intersection. These outputs are machine_predicted_unvalidated and are not human labels.
Distilled classifier scores by category (both heads)
| Category | Codex | Gemma |
|---|---|---|
| Metaresearch | 0.000 | 0.001 |
| Meta-epidemiology (narrow) | 0.000 | 0.000 |
| Meta-epidemiology (broad) | 0.000 | 0.000 |
| Bibliometrics | 0.000 | 0.000 |
| Science and technology studies | 0.000 | 0.000 |
| Scholarly communication | 0.001 | 0.000 |
| Open science | 0.000 | 0.000 |
| Research integrity | 0.000 | 0.000 |
| Insufficient payload (model declined to judge) | 0.001 | 0.000 |
Machine scores (provisional)
The two teacher heads of the student model, read on this work. A score orders the frame for review; it never asserts a category, and the validation status ships verbatim with every row.
Baseline scores from an immature model (maturity gate not passed, 7 training rounds). Scores rank; they never assert a category.
score_only:v0-immature-baseline · verbatim from the scoring run: score_only means the number may rank works, and no category label ships from itClassification
machine, unvalidatedMachine predicted; a candidate call from one source (direct Gemma or distilled Codex), not a consensus.
How this classification was reached, model by model and score by score, is at the end of the page under "How this classification was reached".