In most eukaryotes, homologous chromosomes (homologs) are closely apposed during the prophase of the first meiotic division by a ladderlike proteinaceous structure, the synaptonemal complex (SC) [Fawcett, J Biophys Biochem Cytol 2:403-406, 1956; Moses, J Biophys Biochem Cytol 2: [215][216][217][218] 1956]. SCs consist of two proteinaceous axes, which each support the two sister chromatids of one homolog, and numerous transverse filaments (TFs), which connect the two axes. Organisms that assemble SCs perform meiotic recombination in the context of these structures. Although much information has accumulated about the composition of SCs and the pathways of meiotic crossing over, several questions remain about the role of SCs in meiosis, in particular, about the role of the TFs. In this review, we focus on possible role(s) of TFs. The interest in TF functions received new impulses from the recent characterization of TF-deficient mutants in a number of species. Intriguingly, the phenotypes of these mutants are very different, and a variety of TF functions appear to be hidden behind a façade of morphological conservation. However, in all TFdeficient mutants a specific class of crossovers that display interference is affected. TFs appear to create suitable preconditions for the formation of these crossovers in most species, but are most likely not directly involved in the interference process itself. Furthermore, TFs are important for full-length homolog alignment. (Fig. 1).
Assembly and composition of synaptonemal complexesCohesins, which are proteins that mediate cohesion between sister chromatids (reviewed by Nasmyth 2005), are important for AE assembly (Klein et al. 1999;Pelttari et al. 2001). During S-phase, both in the mitotic cycle and in premeiotic S-phase, cohesins are installed in such a way that they hold the newly synthesized sister chromatids together. In the mitotic cycle, all sister chromatid cohesion is lost at the metaphase-to-anaphase transition, so that sister chromatids can disjoin. In meiosis, two chromosome segregations follow a single round of DNA-replication, and this is possible because cohesion is released in two steps (reviewed by Nasmyth 2001Nasmyth , 2005.Cohesins not only provide sister chromatid cohesion, but also participate in homologous recombination, both in the mitotic cycle and in meiosis. In mitosis, they enhance sister chromatid-based recombinational repair. In meiosis, their role is modified in such a way that homologous recombination occurs preferentially between chromatids of homologs, in most species, mainly or exclusively by a TF-dependent pathway of crossing over (reviewed by