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. Author manuscript; available in PMC: 2016 Apr 1.
Published in final edited form as: Biol Chem. 2015 Apr 1;396(4):295–310. doi: 10.1515/hsz-2014-0267

Table 1.

List of KASH and SUN protein associations

Species Cytoplasmic Binding proteins KASH proteins SUN proteins Nuclear Binding proteins Functions Methods
S. cerevisiae Bbp1 Mps2 Mps3 SPB integrity Mps2 & Bbp1: coIP, GST pull down (Schramm et al., 2000)
Mps2 & Mps3: IF, Y2H (Friederichs et al., 2011)
Spc24 Mps2 Chromosome segregation Mps2 & Spc24: coIP (Le Masson et al., 2002)
Actin Csm4 Mps3 Ndj1 Meiotic telomere movement

Meiotic telomere tethering
Actin & Csm4: mutant animal study (Conrad et al., 2008; Kosaka et al., 2008; Wanat et al., 2008)
Csm4 & Mps3: coIP (Conrad et al., 2008)
Mps3 & Ndj1: coIP (Conrad et al., 2008; Kosaka et al., 2008)
Mps3 Sir4, ribosome biogenesis factors Telomere tethering
Protection of telomeres from transcription
Mps3 & Sir4: coIP (Bupp et al., 2007)

Mps3 & Ribosome biogenesis factors: coIP (Horigome et al., 2011), Y2H (Horigome et al., 2011)
Mps3 Telomerase, Ku70, Ku80 Telomere tethering
Protection of telomeres from transcription
Mps3 & telomerase: IF of reporter (Schober et al., 2009), GST pull down (Antoniacci et al., 2007)
Mps4 & K70/Ku80: IF of reporter (Antoniacci et al., 2004), mutant animal study (Oza et al., 2009)
Mps3 Replication factor C, Htz1 DNA damage response Mps3 & Replication factor C, Htz1: CoIP, in vitro coupling (Haas et al., 2012)
Mps3 Eco1/Ctf7 Chromosome tethering
Sister chromatin cohesion
Mps3 & Eco1/Ctf7: GST pull down, coIP (Antoniacci et al., 2004)
S. pombe Dynein (Dlc1) Kms1 Sad1 Bqt1, Bqt2 Meiotic telomere movement

Meiotic telomere tethering
Dlc1 & Kms1: Y2H (Miki et al., 2004), IF (Yoshida et al., 2013)
Kms1 & Sad1: Y2H (Miki et al., 2004)
Sad1 & Bqt1/2: IF, coIP, mutant animal study (Chikashige et al., 2006)
Cut12, Pcp1 Kms2 Sad1 SPB integrity and tethering Cut12 & Kms2: IF, GST pull down (Walde and King, 2014)
Kms2 & Sad1: IF (King et al., 2008)
Sad1 Csi Centromere tethering Sad1 & Csi1: coIP (Hou et al., 2012)
D. melanogaster Yuri Gagarin, dynein, dynactin Spag4 Centriolar coupling during spermatogenesis Yuri Gagarin, dynein, dynactin & Spag4: IF (Kracklauer et al., 2010)
Microtubule Klarsicht Klaroid Lam Dm0 Nuclear migration during eye development Microtubule & Klarsich: IF (Fischer et al., 2004; Patterson et al., 2004)
Klarisicht & Klaroid: mutant animal study (Kracklauer et al., 2007)
Klaroid & Lam Dm0: IF (Patterson et al., 2004)
Actin MSP-300 Klaroid Lam Dm0 Nuclear anchorage in muscle fiber Actin & MSP-300: IF, in vitro coupling (Volk, 1992)
MSP-300 & Klaroid: mutant animal study (Technau and Roth, 2008)
Klar MSP-300 Klaroid Lam Dm0 Microtubule coupling MSP-300 & Klar: coIP (Elhanany-Tamir et al., 2012)
C. elegans Actin Anc-1 Unc-84 Ce-Lamin Nuclear anchorage Actin & Anc-1: in vitro coupling (Starr and Han, 2002)
Anc-1 & Unc-84: mutant animal study (Starr and Han, 2002)
Unc-84 & Ce Lamin: mutant animal study (Lee et al., 2002)
Rpm1 Anc-1 Positive regulation of Bar-1 during neuronal development Rpm1 &Anc1: coIP (Tulgren et al., 2014)
Dynein (DLC-1/BICD-1/Nud-2) Unc-83 Unc-84 Ce-Lamin Nuclear migration of hypodermal cells Dynein & Unc-83: GST pull down, coIP (Fridolfsson et al., 2010)
Unc-83 & Unc-84: mutant animal study (Starr et al., 2001)
Kinesin (KLC-2) Unc-83 Unc-84 Ce-Lamin Nuclear migration of hypodermal cells Kinesin & Unc83: GST pull down, coIP, IF (Meyerzon et al., 2009)
Zyg-12A Zyg-12 Sun1/MTF1 Centrosome nucleus coupling Zyg-12A & Zyg12: coIP (Malone et al., 2003)
Zyg12 & Sun1/MTF: mutant animal study (Malone et al., 2003)
Dynein (DLI-1/Lis-1/Arp-1/DHC1) Zyg-12 Sun1/MTF1 Him-8, Zim1, Zim2, Zim3 Meiotic chromosome movement

Meiotic chromosome tethering
Dynein & Zyg12: IF, mutant animal study (Malone et al., 2003; Sato et al., 2009; Zhou et al., 2009)
Sun1 & Him-8, Zim-1,-2,-3: IF (Sato et al., 2009)
KDP-1 Sun1/MTF1 Cell cycle progression Kdp1 & Sun1/MTF1: Y2H, mutant animal study (McGee et al., 2009)
Sun1/MTF1 CED Recruit CED to nucleus Sun1 & CED: in vitro coupling (Tzur et al., 2006a)
D. rerio LRMP Pronuclear fusion
Centrosome nucleus coupling
Binding partners unknown (Lindeman and Pelegri, 2012)
Vertebrate Actin Nesprin-1 Sun1/Sun2 Lamin A/C Nuclear anchorage at the NMJ Actin & Nesprin-1: in vitro coupling(Padmakumar et al., 2004), IP (Nikolova-Krstevski et al., 2011)
Nesprins & Sun1,2: coIP (Crisp et al., 2006; Haque et al., 2006)
Sun1,2 & Atype lamins: GST pull down (Crisp et al., 2006), coIP (Haque et al., 2006)
Dynein, dynactin Nesprin-1 Sun1/Sun2 Lamin A/C Nuclear migration during neuronal development Dynein & Nesprin-1: IF, IP (Yu et al., 2011; Zhang et al., 2009)
Nesprin3 Nesprin-1 Sun1/Sun2 Lamin A/C Regulation of nuclear size Nepsrin1 & Nesprin3: GST, His-tag pull down, Y2H (Taranum et al., 2012)
Nesprin-1 Sun3 Formation of acrosome Nesprin-1 & Sun3: IF (Gob et al., 2010)
Actin Nesprin-2 Sun1/Sun2 Lamin A/C Nuclear migration during neuronal development Actin & Nesprin-2: in vitro coupling (Zhen et al., 2002)
Nesprin-2 & Sun1/2: coIP (Crisp et al., 2006; Haque et al., 2006)
Dynein, dynactin Nesprin-2 Sun1/Sun2 Lamin A/C Nuclear migration during neuronal development Dynein & Nesprin-2: IP, IF (Yu et al., 2011; Zhang et al., 2009)
Kinesin Nesprin-2 Sun1/Sun2 Lamin A/C Nuclear migration during retinal development Kinesin & Nesprin-2: GST pull down (Schneider et al., 2011), IP (Yu et al., 2011)
Plectin Nesprin-3 Sun1/Sun2 Lamin A/C Intermediate filament coupling Plectin & Nesprin3: coIP, IF (Ketema et al., 2007; Wilhelmsen et al., 2005)
Nesprin3 & Sun1/2: coIP, IF(Ketema et al., 2007)
Nesprin-3 Sun1η Formation of sperm heard Nesprin3 & Sun1η: IF (Gob et al., 2010)
KLC-1/kif5b Nepsrin-4 Sun1/Sun2 Lamin A/C Nuclear positioning of secretory epithelia cells
Nuclear positioning of inner ear cells
KLC-1 & Nesprin4: coIP, IF (Roux et al., 2009), GST pull down (Wang et al., 2013)

Nesprin4 & Sun1/2: IF (Roux et al., 2009)
Dynein, Dynactin KASH5 Sun1 Meiotic telomere movement Dynein & KASH5: IP (Morimoto et al., 2012), coIP, IF (Horn et al., 2013b)
KASH5 & Sun1: IF (Horn et al., 2013b)
IP3R LRMP Sun1/Sun2 Lamin A/C IP3R recruitment to the NE IP3R & LRMP: coIP, IF (Shindo et al., 2010)
LRMP & Sun1: IF (Horn et al., 2013b)
Nesprin-1/-2 LaminA/C Emerin Nesprin-1/Nesprin-2 & LaminA/C: GST pull down (Mislow et al., 2002), coIP, IP (Zhang et al., 2005)
Nesprin-1/Nesprin-2 & Emerin: in vitro coupling (Mislow et al., 2002), GST pull down (Wheeler et al., 2007), coIP, IP (Zhang et al., 2005)
TorsinA Nesprin-1/-2/-3 TorsinA & Nesprins: coIP (Nery et al., 2008)
Sun1 DNAPK, Ku70, Ku80, NAT10 DNA damage response Sun1 & DNPK: coIP (Lei et al., 2012)
Sun1 & NAT10:coIP (Chi et al., 2007)
Sun1/Sun2 SAMP1 Cell migration Sun2 & SAMP1: IF, coIP (Borrego-Pinto et al., 2012)
Sun1 & SAMP1: MCLIP(Jafferali et al., 2014)
RanGAP AtWips AtSun1/2 LINC Recruitment of RanGAP RanGAP & AtWips: IP, IF (Meier et al., 2010; Xu et al., 2007)
AtWips & AtSuns: coIP, FRAP (Zhou et al., 2012a)
AtSun1/2 & LINC: FRET (Graumann, 2014)
A. thaliana WIT1, 2 AtWips AtSun1/2 Actin coupling AtWit1,2 & AtWips: IP (Zhao et al., 2008)
WIT1, 2 AtWips AtSun1/2 Actin coupling AtWit1,2 & AtWips: IP (Zhao et al., 2008)

coIP: co-immunoprecipitation, IF: immunofluorescence, Y2H: yeasttwo-hybrid, IP: immunoprecipitation, MCLIP: Membrane protein cross-link immunoprecipitation