Torsions-Driven Root Helical Growth, Waving And Skewing In Arabidopsis

Helical growth broadly exists in immobile plants to support their limited movement, and Arabidopsis seedling root exhibiting natural left-handedness helical growth is considered as a simplified model for investigating this interesting behavior. Efforts have been made for understanding the mechanism of root helical growth and consequent root waving and skewing on tilted and impenetrable surface, and several models have been established. Here, previous reports are reviewed and a straightforward torsions-driven mechanism has been emphasized, and additional experiments have been performed to fill up the gaps of this theory in our study.This study implies that, torsions originating from handedness of both cortical microtubules and cellulose microfibrils play central role in root handed helical growth. Different from torsions directly provided by handed assembled cortical microtubules, torsions originating from right-handed assembled cellulose microfibrils are relaxed by their cross-linking with pectin within cell wall, but only exhibited when their cross-linking is interrupted due to damaged cell wall integrity. To topologically relax these torsions, supercoils of cortical microtubules and/or cellulose microfibrils exhibiting as oblique alignments are formed in root cells, which alter the orientation of root cell files and generate handed helical roots. Working together with gravitropic response, relaxation of torsions originating from helical roots drives roots to elongate with handedness, which therefore produces waved and skewed roots on tilted and impenetrable surface.

Dinitroaniline Herbicide Resistance and Mechanisms in Weeds

Dinitroanilines are microtubule inhibitors, targeting tubulin proteins in plants and protists. Dinitroaniline herbicides, such as trifluralin, pendimethalin and oryzalin, have been used as pre-emergence herbicides for weed control for decades. With widespread resistance to post-emergence herbicides in weeds, the use of pre-emergence herbicides such as dinitroanilines has increased, in part, due to relatively slow evolution of resistance in weeds to these herbicides. Target-site resistance (TSR) to dinitroaniline herbicides due to point mutations in α-tubulin genes has been confirmed in a few weedy plant species (e.g., Eleusine indica, Setaria viridis, and recently in Lolium rigidum). Of particular interest is the resistance mutation Arg-243-Met identified from dinitroaniline-resistant L. rigidum that causes helical growth when plants are homozygous for the mutation. The recessive nature of the TSR, plus possible fitness cost for some resistance mutations, likely slows resistance evolution. Furthermore, non-target-site resistance (NTSR) to dinitroanilines has been rarely reported and only confirmed in Lolium rigidum due to enhanced herbicide metabolism (metabolic resistance). A cytochrome P450 gene (CYP81A10) has been recently identified in L. rigidum that confers resistance to trifluralin. Moreover, TSR and NTSR have been shown to co-exist in the same weedy species, population, and plant. The implication of knowledge and information on TSR and NTSR in management of dinitroaniline resistance is discussed.

Helical growth during the phototropic response, avoidance response, and in stiff mutants of Phycomyces blakesleeanus

The sporangiophores of Phycomyces blakesleeanus have been used as a model system to study sensory transduction, helical growth, and to establish global biophysical equations for expansive growth of walled cells. More recently, local statistical biophysical models of the cell wall are being constructed to better understand the molecular underpinnings of helical growth and its behavior during the many growth responses of the sporangiophores to sensory stimuli. Previous experimental and theoretical findings guide the development of these local models. Future development requires an investigation of explicit and implicit assumptions made in the prior research. Here, experiments are conducted to test three assumptions made in prior research, that (a) elongation rate, (b) rotation rate, and (c) helical growth steepness, R, of the sporangiophore remain constant during the phototropic response (bending toward unilateral light) and the avoidance response (bending away from solid barriers). The experimental results reveal that all three assumptions are incorrect for the phototropic response and probably incorrect for the avoidance response but the results are less conclusive. Generally, the experimental results indicate that the elongation and rotation rates increase during these responses, as does R, indicating that the helical growth steepness become flatter. The implications of these findings on prior research, the “fibril reorientation and slippage” hypothesis, global biophysical equations, and local statistical biophysical models are discussed.

Mechanistic Insights into Plant Chiral Growth

The latent left–right asymmetry (chirality) of vascular plants is best witnessed as a helical elongation of cylindrical organs in climbing plants. Interestingly, helical handedness is usually fixed in given species, suggesting genetic control of chirality. Arabidopsis thaliana, a small mustard plant, normally does not twist but can be mutated to exhibit helical growth in elongating organs. Genetic, molecular and cell biological analyses of these twisting mutants are providing mechanistic insights into the left–right handedness as well as how potential organ skewing is suppressed in most plants. Growth direction of elongating plant cells is determined by alignment of cellulose microfibrils in cell walls, which is guided by cortical microtubules localized just beneath the plasma membrane. Mutations in tubulins and regulators of microtubule assembly or organization give rise to helical arrangements of cortical microtubule arrays in Arabidopsis cells and cause helical growth of fixed handedness in axial organs such as roots and stems. Whether tubulins are assembled into a microtubule composed of straight or tilted protofilaments might determine straight or twisting growth. Mechanistic understanding of helical plant growth will provide a paradigm for connecting protein filament structure to cellular organization.

The loss of function of HEL, which encodes a cellulose synthase interactive protein, causes helical and vine-like growth of tomato

Helical growth is an economical way for plant to obtain resources. The classic microtubule–microfibril alignment model of Arabidopsis helical growth involves restriction of the appropriate orientation of cellulose microfibrils appropriately in the cell walls. However, the molecular mechanism underlying tomato helical growth remains unknown. Here, we identified a spontaneous tomato helical (hel) mutant with right-handed helical cotyledons and petals but left-handed helical stems and true leaves. Genetic analysis revealed that the hel phenotype was controlled by a single recessive gene. Using map-based cloning, we cloned the HEL gene, which encodes a cellulose interacting protein homologous to CSI1 of Arabidopsis. We identified a 27 bp fragment replacement that generated a premature stop codon. Transgenic experiments showed that the helical growth phenotype could be restored by the allele of this gene from wild-type Pyriforme. In contrast, the knockout mutation of HEL in Pyriforme via CRISPR/Cas9 resulted in helical growth. These findings shed light on the molecular control of the helical growth of tomato.

Coupling Large Eddies and Waves in Turbulence: Case Study of Magnetic Helicity at the Ion Inertial Scale

In turbulence, for neutral or conducting fluids, a large ratio of scales is excited because of the possible occurrence of inverse cascades to large, global scales together with direct cascades to small, dissipative scales, as observed in the atmosphere and oceans, or in the solar environment. In this context, using direct numerical simulations with forcing, we analyze scale dynamics in the presence of magnetic fields with a generalized Ohm’s law including a Hall current. The ion inertial length ϵ H serves as the control parameter at fixed Reynolds number. Both the magnetic and generalized helicity—invariants in the ideal case—grow linearly with time, as expected from classical arguments. The cross-correlation between the velocity and magnetic field grows as well, more so in relative terms for a stronger Hall current. We find that the helical growth rates vary exponentially with ϵ H , provided the ion inertial scale resides within the inverse cascade range. These exponential variations are recovered phenomenologically using simple scaling arguments. They are directly linked to the wavenumber power-law dependence of generalized and magnetic helicity, ∼ k − 2 , in their inverse ranges. This illustrates and confirms the important role of the interplay between large and small scales in the dynamics of turbulent flows.