The meiosis-specific HOP1 gene is important both for crossing over between homologs and for production of viable spores. hop1 diploids fail to assemble synaptonemal complex (SC), which normally provides the framework for meiotic synapsis. Immunochemical methods have shown that the 70-kDa HOP1 product is a component of the SC. To assess its molecular function, we have purified Hop1 protein to homogeneity and shown that it forms dimers and higher oligomers in solution. Consistent with the zinc-finger motif in its sequence, the purified protein contained about 1 mol equivalent of zinc whereas mutant protein lacking a conserved cysteine within this motif did not. Electrophoretic gel mobility shift assays with different forms of M13 DNA showed that Hop1 binds more readily to linear duplex DNA and negatively superhelical DNA than to nicked circular duplex DNA and even more weakly to single-stranded DNA. Linear duplex DNA binding was enhanced by the addition of Zn 2؉ , was stronger for longer DNA fragments, and was saturable to about 55 bp/protein monomer. Competitive inhibition of this binding by added oligonucleotides suggests preferential affinity for G-rich sequences and weaker binding to poly(dA-dT). Nuclear extracts of meiotic cells caused exonucleolytic degradation of linear duplex DNA if the extracts were prepared from hop1 mutants; addition of purified Hop1 conferred protection against this degradation. These findings suggest that Hop1 acts in meiotic synapsis by binding to sites of double-strand break formation and helping to mediate their processing in the pathway to meiotic recombination.Meiosis is a crucial step in the cycle of sexual reproduction, since it reduces the chromosome complement to haploidy in preparation for fertilization. A single round of DNA replication is followed by two successive rounds of chromosome segregation to produce four haploid products. During the first division, the centromeres of homologous chromosomes are translocated to opposite poles of the meiotic spindle while sister centromeres remain associated with one another. The fidelity of this division depends on crossing over between the homologs, because the chiasmata thus formed provide for cohesion between the homologs as they become aligned on the metaphase plate. In many organisms, crossing over depends in turn on the elaboration of the synaptonemal complex (SC), which joins the homologs along their length during the period when meiotic recombination takes place (23,32,36). Cytologically, SC is seen as a tripartite structure, consisting of a central element flanked by two lateral elements that lie about 100 nm apart and are interconnected by transverse elements (43). It has been argued persuasively that only those recombination events that occur within the context of the SC generate stable chiasmata that are capable of facilitating proper disjunction (2, 12, 23).The yeast Saccharomyces cerevisiae has served as an instructive model for genetic dissection of key mechanisms in meiosis. Combined genetic and cytological analyses ...