and so on. After each endosymbiosis, the partners eventually eliminated unproductive duplication of genetic functions by re-arranging their genomes, a process which sometimes involved transfer of genes between them. Another hypothesis proposes that mitochondria were originally sulfur- or hydrogen-metabolising endosymbionts, and became oxygen-consumers later. On the other hand, mitochondria might have been part of eukaryotes' original equipment. There is a debate about when eukaryotes first appeared: the presence of steranes in Australian shales may indicate eukaryotes at 2.7 Ga; however, an analysis in 2008 concluded that these chemicals infiltrated the rocks less than 2.2 Ga and prove nothing about the origins of eukaryotes. Fossils of the algae Grypania have been reported in 1.85 billion-year-old rocks (originally dated to 2.1 Ga but later revised), indicating that eukaryotes with organelles had already evolved. A diverse collection of fossil algae were found in rocks dated between 1.5 and 1.4 Ga. The earliest known fossils of fungi date from 1.43 Ga. Plastids Plastids, the superclass of organelles of which chloroplasts are the best-known exemplar, are thought to have originated from endosymbiotic cyanobacteria. The symbiosis evolved around 1.5 Ga and enabled eukaryotes to carry out oxygenic photosynthesis. Three evolutionary lineages of photosynthetic plastids have since emerged: chloroplasts in green algae and plants, rhodoplasts in red algae and cyanelles in the glaucophytes. Not long after this primary endosymbiosis of plastids, rhodoplasts and chloroplasts were passed down to other bikonts, establishing an eukaryotic assemblage of phytoplankton by the end of the Neoproterozoic Eon. Sexual reproduction and multicellular organisms Evolution of sexual reproduction The defining characteristics of sexual reproduction in eukaryotes are meiosis and fertilization, resulting in genetic recombination, giving offspring 50% of their genes from each parent. By contrast, in asexual reproduction there is no recombination, but occasional horizontal gene transfer. Bacteria also exchange DNA by bacterial conjugation, enabling the spread of resistance to antibiotics and other toxins, and the ability to utilize new metabolites. However, conjugation is not a means of reproduction, and is not limited to members of the same species – there are cases where bacteria transfer DNA to plants and animals. On the other hand, bacterial transformation is clearly an adaptation for transfer of DNA between bacteria of the same species. This is a complex process involving the products of numerous bacterial genes and can be regarded as a bacterial form of sex. This process occurs naturally in at least 67 prokaryotic species