The determination of the structure of DNA by Watson and Crick in 1953 is often said to mark the birth of modern molecular biology. The DNA is a large molecule composed of four basic units called nucleotides. Each nucleotide contains phosphate, sugar and one of the four bases: Adenine, Guanine, Cytosine and Thymine (usually denoted as A,G,C and T). The structure of DNA is described as a double helix. Each helix, or strand, is a polymer of nucleotides chained together by phosphodiester bonds. The two helices are held together by hydrogen bonds. These bonds are formed by pairs of bases, with each base pair consisting of one purine base (A or G) and one pyrimidine base (C or T), paired according to the following rule: G pairs with C, and A pairs with T. The DNA molecule is directional, due to the assymetrical structure of the sugars which constitute the skeleton of the molecule. Each sugar is connected to the strand upstream (i.e. before it in the chain) in its fifth carbon and to the strand downstream (i.e. after it) in its third carbon. Therefore we say that the DNA strand goes from 5' (read five prime) to 3' (read three prime). The directions of the two complementary DNA strands are reversed to one another. The total length of the human DNA is estimated to be base pairs (abbreviated bp). The amount of DNA varies between different organisms. The organism Amoeba dubia (a single celled organism), for example, has more than 200 times the DNA than human.