In many morphogenetic events in Drosophila and other organisms, non-centrosomal MTs connect local cell-shape changes to distant positions through specific adaptors (Lee and Harland, 2007; Lee et al., 2007; Yano et al., 2021). Many tissues orient MTs with minus ends toward the apical compartment and the plus ends toward the basal, and this organization is held through MT-binding adaptors (Fig. 2A). One of the very versatile MT regulators is Shot, which depending on the context, gets recruited basally, apically, or in both compartments, and depending on its binding partners, it can interact with MT plus or minus ends. Shot recruitment at MT minus ends is mediated by Patronin, and its plus end localization is mediated by EB1 (Applewhite et al., 2010; Booth et al., 2014; Ghislain et al., 2021; Goodwin and Vale, 2010; Lee et al., 2016; Mimori-Kiyosue et al., 2000; Molines et al., 2018; Nashchekin et al., 2016). Despite the different scenarios in which Shot can regulate MTs and their interaction with apicobasal compartments, its contribution to a given process can vary drastically. For instance, in late pupal wing development, MTs run along the apicobasal axis and they stabilise adhesion to the apical and basal ECMs. Shot is found at both ends, as it is in terminal cells. However, loss of Shot has no effect on MT architecture or epithelial morphology. Instead, in this tissue, MTs are stabilised through their interactions with Pio in the apical membrane and ILK and integrins in the basal domain (Akhtar and Streuli, 2013; Bökel et al., 2005). These results contrast with subcellular tube formation, where Shot seems to contribute more than Pio in mediating apicobasal interactions (Jaźwińska et al., 2003; Ricolo and Araujo, 2020). Shot and Patronin organise MTs at the apical compartment in a range of processes, like in the formation of apical actin-based microvilli of follicle cells and in salivary glands, where they are required for proper tissue invagination (Booth et al., 2014; Ghislain et al., 2021; Khanal et al., 2016; Röper, 2012). In these models, MT apical anchoring is required to sustain novel architectures, but whether they interact with the basal plasma membrane is still unknown.

There are processes that use MT apical anchoring to drive cell-shape changes but do not seem to depend on Shot at all. This is seen for instance in dorsal fold formation during early development, where instead, Patronin organises an MT apical cortex that is required to initiate fold formation. In this process, instead of a transversal array, MTs form an apical dome that allows proper tissue folding (Takeda et al., 2018). Similarly, a morphogenetic furrow regulates proper photoreceptor organisation in the eye disc (Ready et al., 1976). This furrow is formed through apical recruitment of actomyosin which induces apical constriction, and apicobasal MTs are also required for the shape change. Loss of integrins affects MT organisation and furrow formation, but the elements that mediate MT anchoring to the apical and basal compartments have not been reported, and at least involvement of ILK has been ruled out (Corrigall et al., 2007; Fernandes et al., 2014).