title: Nitrogen fixation in eukaryotes--new models for symbiosis.
abstract: BACKGROUND
  Nitrogen, a component of many bio-molecules, is essential for growth and development of all organisms. Most nitrogen exists in the atmosphere, and utilisation of this source is important as a means of avoiding nitrogen starvation. However, the ability to fix atmospheric nitrogen via the nitrogenase enzyme complex is restricted to some bacteria. Eukaryotic organisms are only able to obtain fixed nitrogen through their symbiotic interactions with nitrogen-fixing prokaryotes. These symbioses involve a variety of host organisms, including animals, plants, fungi and protists.
  
  
  RESULTS
  We have compared the morphological, physiological and molecular characteristics of nitrogen fixing symbiotic associations of bacteria and their diverse hosts. Special features of the interaction, e.g. vertical transmission of symbionts, grade of dependency of partners and physiological modifications have been considered in terms of extent of co-evolution and adaptation. Our findings are that, despite many adaptations enabling a beneficial partnership, most symbioses for molecular nitrogen fixation involve facultative interactions. However, some interactions, among them endosymbioses between cyanobacteria and diatoms, show characteristics that reveal a more obligate status of co-evolution.
  
  
  CONCLUSIONS
  Our review emphasises that molecular nitrogen fixation, a driving force for interactions and co-evolution of different species, is a widespread phenomenon involving many different organisms and ecosystems. The diverse grades of symbioses, ranging from loose associations to highly specific intracellular interactions, might themselves reflect the range of potential evolutionary fates for symbiotic partnerships. These include the extreme evolutionary modifications and adaptations that have accompanied the formation of organelles in eukaryotic cells: plastids and mitochondria. However, age and extensive adaptation of plastids and mitochondria complicate the investigation of processes involved in the transition of symbionts to organelles. Extant lineages of symbiotic associations for nitrogen fixation show diverse grades of adaptation and co-evolution, thereby representing different stages of symbiont-host interaction. In particular cyanobacterial associations with protists, like the Rhopalodia gibba-spheroid body symbiosis, could serve as important model systems for the investigation of the complex mechanisms underlying organelle evolution.
authors.affiliation: Department of Cell Biology, Philipps-University Marburg, Marburg, Germany. kneip@mpiib-berlin.mpg.de
authors.forename: Christoph
authors.initials: C
authors.lastname: Kneip
authors.forename: Peter
authors.initials: P
authors.lastname: Lockhart
authors.forename: Christine
authors.initials: C
authors.lastname: Voss
authors.forename: Uwe-G
authors.initials: UG
authors.lastname: Maier
mesh_terms: D000818:Animals; D001419:Bacteria / Q000235:genetics* / Q000378:metabolism; D017377:Diatoms / Q000235:genetics / Q000378:metabolism / Q000502:physiology; D005658:Fungi / Q000235:genetics / Q000378:metabolism / Q000502:physiology; D007448:Invertebrates / Q000235:genetics / Q000382:microbiology / Q000502:physiology; D008954:Models, Biological; D009586:Nitrogen Fixation / Q000235:genetics*; D010802:Phylogeny; D010944:Plants / Q000235:genetics / Q000378:metabolism; D013559:Symbiosis