number (n) is the amount of β strands in the porin, and barrel radius (R) refers to the radius of the opening of the porin. These parameters are related via the following formulas: and, Using these formulas, the structure of a porin can be determined by knowing only a few of the available parameters. While the structure of many porins have been determined using X-ray crystallography, the alternative method of sequencing protein primary structure may also be used instead. Cellular roles Porins are water-filled pores and channels found in the membranes of bacteria and eukaryotes. Porin-like channels have also been discovered in archaea. Note that the term "nucleoporin" refers to unrelated proteins that facilitate transport through nuclear pores in the nuclear envelope. Porins are primarily involved in passively transporting hydrophilic molecules of various sizes and charges across the membrane. For survival, certain required nutrients and substrates must be transported into the cells. Likewise, toxins and wastes must be transported out to avoid toxic accumulation. Additionally, porins can regulate permeability and prevent lysis by limiting the entry of detergents into the cell. Two types of porins exist to transport different materials– general and selective. General porins have no substrate specificities, though some exhibit slight preferences for anions or cations. Selective porins are smaller than general porins, and have specificities for chemical species. These specificities are determined by the threshold sizes of the porins, and the amino acid residues lining them. In gram-negative bacteria, the inner membrane is the major permeability barrier. The outer membrane is more permeable to hydrophilic substances, due to the presence of porins. Porins have threshold sizes of transportable molecules that depend on the type of bacteria and porin. Generally, only substances less than 600 Daltons in size can diffuse through. Diversity Porins were first discovered in gram-negative bacteria, but gram-positive bacteria with both types of porins have been found. They exhibit similar transport functions but have a more limited variety of porins, compared to the distribution found in gram-negative bacteria. Gram-positive bacteria lack outer membranes, so these porin channels are instead bound to specific lipids within the cell walls. Porins are also found in eukaryotes, specifically in the outer membranes of mitochondria and chloroplasts. The organelles contain general porins that are structurally and functionally similar to bacterial ones. These similarities have supported the Endosymbiotic theory, through which eukaryotic organelles arose from gram-negative bacteria. However, eukaryotic porins exhibit