Table 1.

Changes in nuclear shape, nuclear size, chromatin texture, and nuclear protein expression across different cancer types.

Cancer Type	Changes in nuclear shape	Changes in nuclear size	Changes in chromatin/nucleoli	Changes to nuclear proteins
Bladder cancer	Enlarged mean nuclear area (Krüger and Müller 1995) High nuclear to cytoplasmic size ratio (Bhatia et al. 2006)	Variation in nuclear size (Bhatia et al. 2006)	Coarse chromatin (Bhatia et al. 2006)
Breast cancer	Variation in nuclear shape (Abdalla et al. 2009; Kashyap et al. 2018)	Enlarged nuclear area, variation in nuclear area (Abdalla et al. 2009; Kashyap et al. 2018)		Reduced levels or loss of lamin A/C (Alhudiri et al. 2019; Capo-Chichi et al. 2011b; Matsumoto et al. 2015) Ratio of mRNA levels for lamin C:lamin A increased in breast cancer samples (Aljada et al. 2016) Loss of SUN1, SUN2, nesprins (Matsumoto et al. 2015) Mutated nesprins 1, 2 (Sjöblom et al. 2006; Stephens et al. 2012) Nup88 overexpression (Agudo et al. 2004; Gould et al. 2002)
Cervical cancer	Irregular nuclear shapes (Papanicolaou and Traut 1941)	Large nuclei (Papanicolaou and Traut 1941)	Speckled chromatin, one or more nucleoli (Papanicolaou and Traut 1941)	Nup88 overexpression (Brustmann and Hager 2009)
Colorectal cancer	Irregular nuclear contours with folds and dents (Fischer 2020) Abnormal morphology (Mulder et al. 1992)	Large nuclei (Fischer 2020)	Coarse chromatin (Fischer 2020; Mulder et al. 1992)	Low levels of lamin A/C expression (Belt et al. 2011; Moss et al. 1999) Abnormal levels of lamin B1/type B lamins (Moss et al. 1999; Alfonso et al. 2008) Aberrant staining of lamins in cytoplasm (Moss et al. 1999) Mutated nesprin 1 (Sjöblom et al. 2006) Nup88 overexpression (Emterling et al. 2003; Knoess et al. 2006)
Gastric cancer				Reduced levels or loss of lamin A/C (Moss et al. 1999; Wu et al. 2009) Reduced lamin B1 levels (Moss et al. 1999) Nup88 overexpression (Gould et al. 2002)
Head and neck cancer	Nuclear contour irregularities and nuclear shape asymmetry(Giardina et al. 1996)	Abnormal variation in cell size (Speight 2007)	Hyperchromatic nuclei, increased number of nucleoli(Speight 2007)
Renal cell carcinoma	Irregular contours, multilobed nuclei (Nativ et al. 1996)	Enlarged nuclei (Nativ et al. 1996)	Prominent and irregular nucleoli (Nativ et al. 1996)	Nup88 overexpression (Gould et al. 2002)
Liver cancer			Abnormal chromatin texture (Atupelage et al. 2014)	Lamin B1 is upregulated (Sun et al. 2010; Lim et al. 2002) Nup88 overexpression (Gould et al. 2002)
Lung cancer	Malleable nuclei with crush artifacts (DeMay 2012) Nuclear pleomorphisms (Petersen et al. 2009)			Reduced expression or absence of B-type lamins in small-cell lung cancers (SCLCs)(Broers et al. 1993) Aberrant, cytoplasmic expression of A-type lamins in non-SCLCs (Broers et al. 1993) Nup88 overexpression (Gould et al. 2002; Zhang et al. 2007b)
Melanoma	Variation in nuclear circularity (Mijovic et al. 2013)	Increased nuclear size (Mijovic et al. 2013)		Reduced or complete loss of lamin A/C (Venables et al. 2001) High levels of lamin A/C (Tilli et al. 2003) Nup88 overexpression (Gould et al. 2002)
Ovarian cancer	Irregular nuclear membrane contours (Liu et al. 1999; Gurley et al. 1994)	Irregular nuclear size (Liu et al. 1999; Gurley et al. 1994)		Heterogeneous expression or total Increased levels of lamin B1 and B2 (Bengtsson et al. 2007) Loss of lamin A/C (Capo-chichi et al. 2011a) Loss of emerin (Capo-chichi et al. 2009) Mutations in nesprins (Doherty et al. 2010) Nup88 overexpression (Martínez et al. 1999; Gould et al. 2002) High levels of lamin A/C (Wang et al. 2009)
Pancreatic cancer	Irregular nuclear shape (Sato et al. 1993; Furuta et al. 1992; Barr Fritcher et al. 2007; Flores et al. 2021)			Overexpression of lamin B1 (Li et al. 2013) Overexpression of Nup88 (Gould et al. 2002)
Papillary thyroid carcinoma	Grooves and long clefts (Fische et al. 2003 ; Rosai et al. 1992)	Variability of nuclear size (Wright and Castles 1987)	Coarse heterochromatin (Fischer et al. 2003)
Prostate cancer	Irregular contours and decreased roundness (Epstein et al. 1984; Diamond et al. 1982)	Enlarged nuclei (Veltri et al. 1994)		Lamin B upregulated (Coradeghini et al. 2006) Nup88 overexpression (Gould et al. 2002)