Table 1.

Summary of components regulating nuclear shape

Organism	Protein	Function		Localization	Associated defect	Reference
S. cerevisiae	Esc1	Establishes Silent Chromatin, involved in telomere clustering and Sir4-mediated silencing		INM (absent beneath the nucleolus)	Overexpression results in nuclear membrane ‘escapades’ formation associated with nucleolus	[60]
S. cerevisiae	Mps3	SPB insertion and duplication		INM, SPB	Overexpression results in membrane proliferation, and a few extensions and protrusions	[88]
S. cerevisiae	Acc1	Acetyl CoA carboxylase, catalyses the first rate-limiting step of de novo fatty acid synthesis		Cytoplasm	INM protrudes into the intermembrane space resulting in ‘islands’ between the INM and ONM in the mutants	[168]
S. cerevisiae	Spo7	Regulatory subunit of Nem1-Spo7 phosphatase involved in phospholipid biosynthesis, dephosphorylates Pah1			Misshaped nuclei with a single ‘flare’ like protrusion that colocalizes with the nucleolus	[137]
S. cerevisiae	Hmg1	Enzyme 3-hydroxy-3-methylglutaryl coenzyme A (HMG-CoA) reductase catalyzing a rate-limiting step of sterol biosynthesis		Cytoplasm, Nuclear associated membranes	Proliferation of closely apposed pair of membrane called ‘karmellae’ with some discontinuities	[169]
S. cerevisiae	Dgk1	CTP-dependent Diacyl glycerol kinase involved in phosphatidic acid biosynthesis		NE, ER	Overexpression results in membrane expansion around nucleus making it irregularly shaped and enlarged	[136]
S. cerevisiae	Pah1 (also referred as Smp2)	Phosphotidate phosphatase that dephosphorylates PA to yield diacylglycerol		Nucleus, NE/ER	Deletion results in enlarged, irregularly shaped nuclei with interconnected lobes	[132]
S. cerevisiae	Nup1, Nup53, Nup60, Nup116 and other Nups	Nucleoporins		NE, NPC	Deletion and/or overexpression is associated with irregular nuclear shape and membrane extensions	[74,104,112,113]
S. cerevisiae	Apq12	Functions in lipid homeostasis along with Brr6, Brl1		NE, ER	Membranous divisions within the NE, invaginations and extensions of NE resulting in abnormal shape	[135]
A. thaliana	CRWN1/4	Crowded nucleus proteins, provide lamin-like stability to plant nucleus		INM	Mutants have nuclei more round and soft than Wild type with irregular nuclear margins	[52,170]
A. thaliana	CRWN2/3	Provide lamin-like stability to plant nucleus		Nucleus, INM	Result in abnormal phenotype only when combined with CRWN1/4	[56,170]
A. thaliana	KAKU4	Required for maintaining spindle shaped nuclei, interacts with CRWN1/4 (kaku- Japanese word for nucleus)		INM	Nuclear membrane invagination and stack formation in smaller and less elongated nuclei	[171]
A. thaliana	SUN1/2	SUN-domain containing components of plant LINC complex required for formation and maintenance of polarized nuclear shape in root hairs		INM	Mutants have nearly round nuclei unlike the highly elongated wild type nuclei in mature root hair	[90]
A. thaliana	WIP12/3	WIPs (WPP domain–interacting proteins) are plant-specific KASH proteins required for maintaining elongated plant nuclei in epidermal cells		INM	Less elongated nuclei	[91]
A. thaliana	SINE1-5, TIK1	Tissue-specific plant KASH domain proteins		INM, NE	Abnormal nuclear morphology and positioning	[92,93]
A. thaliana	Nup136	Nucleoporin		NE, NPC	Mutants display abnormally circular nuclei	[114,115]
Mammals	Lamin A/C	Regulate nuclear shape and rigidity, chromatin attachment and interaction with LINC complex		Nucleus, INM	Altered expression deforms nuclei and changes nuclear stiffness	[43,172,173]
Mammals	Lamin B1/ B2			INM
Mammals	Sun2	SUN protein	Involved in regulation of nuclear shape, movement and positioning	INM	Overexpression deforms nuclei into a flower-shaped lobular structure	[164]
Mammals	Nesprin1	KASH protein		ONM	Misshaped nuclei, low circularity, changes in lamina structure	[89,174]
Mammals	Nup53, Nup93	Nucleoporins		NE, NPC	Abnormally shaped nuclei	[111]