LINCing the nuclear envelope to gametogenesis.

Gametogenesis combines two important features: reduction of the genome content from diploid to haploid by carefully partitioning chromosomes, and the subsequent differentiation into fertilization-competent gametes, which in males is characterized by profound nuclear restructuring. These are quite difficult tasks and require a tight coordination of different cellular mechanisms. Recent studies in the field established a key role for LINC complexes in both meiosis and sperm head formation. LINC complexes comprise SUN and KASH domain proteins that form nuclear envelope (NE) bridges, linking the nucleoskeleton to the cytoskeleton. They are well known for their crucial roles in diverse cellular and developmental processes, such as nuclear positioning and cell polarization. In this review, we highlight key roles ascribed to LINC complexes and to the nucleocytoskeletal connection in gametogenesis. First, we give a short overview about the general features of LINC components and the profound reorganization of the NE in germ cells. We then focus on specific roles of LINC complexes in meiotic chromosome dynamics and their impact on pairing, synapsis, and recombination. Finally, we provide an update of the mechanisms controlling sperm head formation and discuss the role of sperm-specific LINC complexes in nuclear shaping and their relation to specialized cytoskeletal structures that form concurrently with nuclear restructuring and sperm elongation.