In animals, the nuclear lamina is a fibrillar meshwork on the inner surface of the nuclear envelope, composed of coiled-coil lamin proteins and lamin binding membrane proteins. Plants also have a meshwork on the inner surface of the nuclear envelope, but little is known about its composition other than the presence of members of the CROWDED NUCLEI (CRWN) protein family, possible plant lamin analogs. Here, we describe a candidate lamina component, based on two Arabidopsis thaliana mutants (kaku2 and kaku4) with aberrant nuclear morphology. The responsible gene in kaku2 encodes CRWN1, and the responsible gene in kaku4 encodes a plant-specific protein of unknown function (KAKU4) that physically interacts with CRWN1 and its homolog CRWN4. Immunogold labeling revealed that KAKU4 localizes at the inner nuclear membrane. KAKU4 deforms the nuclear envelope in a dose-dependent manner, in association with nuclear membrane invagination and stack formation. The KAKU4-dependent nuclear envelope deformation was enhanced by overaccumulation of CRWN1, although KAKU4 can deform the nuclear envelope even in the absence of CRWN1 and/or CRWN4. Together, these results suggest that plants have evolved a unique lamina-like structure to modulate nuclear shape and size.

The nuclear lamina plays an important role in the regulation of chromatin organization and gene positioning in animals. CROWDED NUCLEI (CRWN) is a strong candidate for the plant nuclear lamina protein in Arabidopsis thaliana but its biological function was largely unknown. Here, we show that CRWNs localize at the nuclear lamina and build the meshwork structure. Fluorescence in situ hybridization and RNA-seq analyses revealed that CRWNs regulate chromatin distribution and gene expression. More than 2000 differentially expressed genes were identified in the crwn1crwn4 double mutant. Copper-associated (CA) genes that form a gene cluster on chromosome 5 were among the downregulated genes in the double mutant exhibiting low tolerance to excess copper. Our analyses showed this low tolerance to copper was associated with the suppression of CA gene expression and that CRWN1 interacts with the CA gene locus, enabling the locus to localize at the nuclear lamina under excess copper conditions.

The linear Boussinesq equation should be used in numerical simulations for distant tsunamis in the Pacific Ocean, because the dispersion effect is not negligible. Its difference equation which can not be expressed by an explicit scheme requires a long CPU time. One of the present authors has introduced a new technique, in which the first term of discretization error in the difference equation of the linear long wave theory was used and controlled to replace the physical dispersion term. This method is applied in the present study. The effect of ocean current on the tsunami propagation is confirmed to be negligible in wave direction and wave height. The 1960 Chilean tsunami is simulated. The Coriolis force has not only the effect on the propagation direction but also the dispersion effect, which is examined by comparing with the computed result without the Coriolis force. Effect of the sea bottom topography is examined in detail in terms of wave energy. About 40% of initial wave energy remains on the continental shelf in the neighborhood of Chile, and the rest is radiated to the Ocean. After scattered and trapped by islands and sea mounts, about 25% of the total energy arrives at Japan. The computed results shows a fairly good agreement with tide records after making correction of the effect of the water depth. For further discussions of the tsunami in shallow seas, simulations should be performed using the shallow-water theory and detailed topography.

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