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KASH (Klarsicht, Anc-1, Syne-1 homology) domain-containing proteins anchor the nucleus to the actin cytoskeleton or to microtubules. KASH proteins thus play pivotal roles in a variety of developmental processes where nuclear positioning is critical. Two KASH proteins have been identified in Drosophila: Muscle-specific protein-300 (Msp-300) and Klarsicht (Klar). Msp-300 anchors nuclei to actin, and has been reported to be essential for positioning of nurse cell nuclei during oogenesis, and thus production of mature ooctyes. Klar is required for positioning of photoreceptor and cone cell nuclei in the developing eye, which is critical for proper eye morphology. Here, we asked whether KASH domain-containing forms of Msp-300 are required for nuclear positioning in the eye, and we found that they are not. Moreover, in the course of this work, we discovered that contrary to previous reports, KASH domain-containing forms of Msp-300 are not required for viability, nor for oogenesis. However, we did find that Msp-300 has a function in egg laying, normally redundant with a function of Klar.

KASH (Klarsicht/ Anc-1/ Syne homology) domain proteins are cytoskeleton-associated proteins localized uniquely to the outer nuclear membrane. Klarsicht is a KASH protein required for nuclear migration in differentiating cells of the Drosophila eye. The C-terminal KASH domain of Klarsicht resides in the perinuclear space, and the cytoplasmic moiety connects to the microtubule organizing center. In C. elegans and vertebrate cells, SUN (Sad1/ UNC-84) domain proteins reside in the inner nuclear membrane and tether KASH proteins to the outer nuclear membrane. Is there a Drosophila SUN protein that performs a similar function, and if so, is it like Klarsicht, obviously essential for nuclear positioning only in the eye? Here, we identify Drosophila Klaroid, a SUN protein that tethers Klarsicht. klaroid loss-of-function mutants are indistinguishable phenotypically from klarsicht mutants. Remarkably, neither gene is essential for Drosophila viability or fertility, and even in klaroid klarsicht double mutants, the only obvious external morphological defect is rough eyes. In addition, we find that klaroid and klarsicht are required for nuclear migration in differentiating neurons and in non-neural cells. Finally, while perinuclear Klaroid is ubiquitous in the eye, Klarsicht expression is limited to differentiating cells and may be part of the trigger for apical nuclear migration.