At fertilization, the highly condensed and transcriptionally inert chromatin of the spermatozoa becomes remodelled into the decondensed and transcriptionally competent chromatin of the male pronucleus. The chromatin initially becomes dispersed and then transiently recondenses into a small mass upon entry into the ooplasm. This morphological change is coincident with and likely dependent on the replacement of the sperm-specific protamines by oocyte-supplied histones and the organization of the chromatin into nucleosomes. The chromatin then extensively decondenses within the male pronucleus and acquires many of the proteins that are associated with the maternal chromatin. Nonetheless, the paternal chromatin manifests distinct characteristics, including transient hyperacetylation of histone H4, increased transcription of endogenous and microinjected genes, and replication-independent demethylation of DNA. Sperm chromatin remodelling is controlled by an oocyte activity that appears during meiotic maturation and disappears approximately 3 h after activation (release from metaphase II arrest), and which requires factors associated with the germinal vesicle of the oocyte. The molecular components of this activity remain largely unknown. In frogs, nucleoplasmin is required to assemble histones H2A and H2B onto the paternal chromatin. Evidence is presented that related proteins may perform similar functions in mammals. Identifying the mechanisms that underlie sperm chromatin remodelling at fertilization may be relevant for understanding reprogramming of somatic cell nuclei after transfer into oocytes.Fertilization achieves two objectives: it re-initiates the cell cycle of the dormant egg, and it produces an embryo that contains one functional copy each of the maternal and paternal genomes. The second objective is ensured through a variety of mechanisms, including the block to polyspermy, the completion of the second meiotic division in the oocyte, and the biochemical remodelling of the sperm chromatin to form the male pronucleus. Remodelling of the paternal chromatin at fertilization is necessary because although the paternal genome is required for embryonic development, the DNA of the mature spermatozoa is compacted and transcriptionally inactive. The process through which this remodelling occurs is reviewed here.Biparental androgenetic and gynogenetic mammalian embryos produced by pronuclear transplantation arrest development at mid-gestation (McGrath and Solter, 1984;Surani et al., 1984). These well-known experiments of McGrath and Solter (1984) and Surani et al.