Cortical microtubule arrays are critical in determining the growth axis of diffusely growing plant cells, and various environmental and physiological factors are known to affect the array organization. Microtubule organization is partly disrupted in the spiral1 mutant of Arabidopsis thaliana, which displays a right-handed helical growth phenotype in rapidly elongating epidermal cells. We show here that mutations in the plasma membrane Na(+)/H(+) antiporter SOS1 and its regulatory kinase SOS2 efficiently suppressed both microtubule disruption and helical growth phenotypes of spiral1, and that sos1 and sos2 roots in the absence of salt stress exhibited altered helical growth response to microtubule-interacting drugs at low doses. Salt stress also altered root growth response to the drugs in wild-type roots. Suppression of helical growth appeared to be specific to spiral1 since other helical growth mutants were not rescued. The effects of sos1 in suppressing spiral1 defects and in causing abnormal drug responses were nullified in the presence of the hkt1 Na(+) influx carrier mutation in roots but not in hypocotyls. These results suggest that cytoplasmic salt imbalance caused by insufficient SOS1 activity compromises cortical microtubule functions in which microtubule-localized SPIRAL1 is specifically involved.
Cells at the elongation zone expand longitudinally to form the straight central axis of plant stems, hypocotyls and roots, and transverse cortical microtubule arrays are generally recognized to be important for the anisotropic growth. Recessive mutations in either of two Arabidopsis thaliana SPIRAL loci, SPR1 or SPR2, reduce anisotropic growth of endodermal and cortical cells in roots and etiolated hypocotyls, and induce right-handed helical growth in epidermal cell files of these organs. spr2 mutants additionally show right-handed twisting in petioles and petals. The spr1spr2 double mutant's phenotype is synergistic, suggesting that SPR1 and SPR2 act on a similar process but in separate pathways in controlling cell elongation. Interestingly, addition of a low dose of either of the microtubule-interacting drugs propyzamide or taxol in the agar medium was found to reduce anisotropic expansion of endodermal and cortical cells at the root elongation zone of wild-type seedlings, resulting in left-handed helical growth. In both spiral mutants, exogenous application of these drugs reverted the direction of the epidermal helix, in a dose-dependent manner, from right-handed to left-handed; propyzamide at 1 microM and taxol at 0.2-0.3 microM effectively suppressed the cell elongation defects of spiral seedlings. The spr1 phenotype is more pronounced at low temperatures and is nearly suppressed at high temperatures. Cortical microtubules in elongating epidermal cells of spr1 roots were arranged in left-handed helical arrays, whereas the highly isotropic cortical cells of etiolated spr1 hypocotyls showed microtubule arrays with irregular orientations. We propose that a microtubule-dependent process and SPR1/SPR2 act antagonistically to control directional cell elongation by preventing elongating cells from potential twisting. Our model may have implicit bearing on the circumnutation mechanism.
Variational Bayesian Multimodal EncephaloGraphy (VBMEG) is a MATLAB toolbox that estimates distributed source currents from magnetoencephalography (MEG)/electroencephalography (EEG) data by integrating functional MRI (fMRI) ( https://vbmeg.atr.jp/ ). VBMEG also estimates whole-brain connectome dynamics using anatomical connectivity derived from a diffusion MRI (dMRI). In this paper, we introduce the VBMEG toolbox and demonstrate its usefulness. By collaborating with VBMEG's tutorial page ( https://vbmeg.atr.jp/docs/v2/static/vbmeg2_tutorial_neuromag.html ), we show its full pipeline using an open dataset recorded by Wakeman and Henson ( 2015 ). We import the MEG data and preprocess them to estimate the source currents. From the estimated source currents, we perform a group analysis and examine the differences of current amplitudes between conditions by controlling the false discovery rate (FDR), which yields results consistent with previous studies. We highlight VBMEG's characteristics by comparing these results with those obtained by other source imaging methods: weighted minimum norm estimate (wMNE), dynamic statistical parametric mapping (dSPM), and linearly constrained minimum variance (LCMV) beamformer. We also estimate source currents from the EEG data and the whole-brain connectome dynamics from the MEG data and dMRI. The observed results indicate the reliability, characteristics, and usefulness of VBMEG.
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