Table 1.

Nuclear envelope structural elements that regulate nuclear morphology.

Protein	Organism/ System	Function	Alteration and nuclear phenotype
Lamin A/C	Vertebrates	Structural support to the nucleus, roles in DNA replication and gene expression	Mutations in LMNA gene cause diseases with misshaped nuclei [34- 40]
Lamin A/C	Neutrophil- differentiated HL-60 cells	As above	Overexpression causes hypolobulated nuclei [30,31]
Lamin B1	Mammals Cortical neurons	As above	Deficiency causes misshaped nuclei, nuclear blebs [32]
Lamin B2	As above	As above	Deficiency leads to elongated nuclei [32]
Lamin B2	Zebrafish embryo	As above	Overexpression causes lobulated nuclei with intranuclear membranes [16]
Lamin B3	Xenopus egg extract	As above	Depletion decreases nuclear size [13] Ectopic addition increases nuclear size [14]
Lamin-like nuclear proteins	Arabidopsis thaliana	Should have similar functions to vertebrate lamina	Deletions of LINC1/2 genes result in smaller and more round nuclei [17]
LBR	Neutrophil- differentiated HL-60 cells	Binds lamins and chromatin	Depletion causes hypolobulated nuclei [30,31]
Lap2β	Xenopus egg extract	Binds lamins and chromatin	Addition of truncated Lap2β causes small scalloped nuclei [19]
LEM4	Caenorhabditis elegans	Interacts with lamina and chromatin	Depletion causes misshapen, multilobed nuclei [58]
AtSun1/2	Arabidopsis thaliana	Components of LINC complex	Depletion causes nuclear rounding [46]
AtWIPs	Arabidopsis thaliana	Plant specific KASH domain proteins, components of LINC complex	Mutations that disrupt LINC interactions cause nuclear rounding [47]
Nesprin2	HaCaT cells	KASH domain protein; binds to actin cytoskeleton and SUN proteins; part of LINC complex	Nes2ΔABD overexpression increases nuclear size; Nes2-mini overexpression decreases nuclear size [21,22]
Pom121	Xenopus egg extract	Nucleoporin, structural component of NPC	Addition of dominant negative fragment blocks NE growth [26]
Nup188	Xenopus egg extract	As above	Depletion increases nuclear size [27]
AtNup136	Arabidopsis thaliana	As above	Overexpression increases nuclear size and nuclear elongation; Depletion decreases nuclear size and causes nuclear rounding [44,45]