horizontal gene transfer, the transmission of DNA (deoxyribonucleic acid) between different genomes. Horizontal gene transfer is known to occur between different species, such as between prokaryotes (organisms whose cells lack a defined nucleus) and eukaryotes (organisms whose cells contain a defined nucleus), and between the three DNA-containing organelles of eukaryotes—the nucleus, the mitochondrion, and the chloroplast. Acquisition of DNA through horizontal gene transfer is distinguished from the transmission of genetic material from parents to offspring during reproduction, which is known as vertical gene transfer.
Horizontal gene transfer is made possible in large part by the existence of mobile genetic elements, such as plasmids (extrachromosomal genetic material), transposons (“jumping genes”), and bacteria-infecting viruses (bacteriophages). These elements are transferred between organisms through different mechanisms, which in prokaryotes include transformation, conjugation, and transduction. In transformation, prokaryotes take up free fragments of DNA, often in the form of plasmids, found in their environment. In conjugation, genetic material is exchanged during a temporary union between two cells, which may entail the transfer of a plasmid or transposon. In transduction, DNA is transmitted from one cell to another via a bacteriophage.
In horizontal gene transfer, newly acquired DNA is incorporated into the genome of the recipient through either recombination or insertion. Recombination essentially is the regrouping of genes, such that native and foreign (new) DNA segments that are homologous are edited and combined. Insertion occurs when the foreign DNA introduced into a cell shares no homology with existing DNA. In this case, the new genetic material is embedded between existing genes in the recipient’s genome.
Compared with prokaryotes, the process of horizontal gene transfer in eukaryotes is much more complex, mainly because acquired DNA must pass through both the outer cell membrane and the nuclear membrane to reach the eukaryote’s genome. Subcellular sorting and signaling pathways play a central role in the transport of DNA to the genome.
Prokaryotes can exchange DNA with eukaryotes, although the mechanisms behind this process are not well understood. Suspected mechanisms include conjugation and endocytosis, such as when a eukaryotic cell engulfs a prokaryotic cell and gathers it into a special membrane-bound vesicle for degradation. It is thought that in rare instances in endocytosis, genes escape from prokaryotes during degradation and are subsequently incorporated into the eukaryote’s genome.
Horizontal gene transfer plays an important role in adaptation and evolution in both prokaryotes and eukaryotes. For example, the transfer of a gene encoding a unique metabolic enzyme from a species of Pasteurella bacteria to the protozoan parasite Trichomonas vaginalis is suspected to have facilitated the latter organism’s adaptation to its animal hosts. Likewise, the exchange of a gene from a human cell to the bacterium Neisseria gonorrhoeae—a transfer that appears to have occurred relatively recently in the bacterium’s evolution—may have enabled the organism to adapt and survive in humans. Scientists have proposed too that the recent evolution of the methylaspartate pathway of metabolism in the halophilic (salt-loving) archaean Haloarcula marismortui originated with the organism’s acquisition of a specialized set of genes via horizontal transfer.