text: Genes of all organisms, except bacteria, consist of short protein-coding regions (exons) interrupted by long sequences (introns). When a gene is expressed, its DNA sequence is copied into a “primary RNA” sequence by the enzyme RNA polymerase. Then the “spliceosome” machinery physically removes the introns from the RNA copy of the gene by the process of splicing, leaving only a contiguously connected series of exons, which becomes messenger RNA (mRNA). This mRNA is now read by the ribosome, which produces the encoded protein. Thus, although introns are not physically removed from a gene, a gene's sequence is read as if introns were not present.
  Exons are usually short, with an average length of about 120 bases (e.g. in human genes). Intron lengths vary widely from 10 to 500,000, but exon lengths have an upper bound of about 600 bases in most eukaryotes. Because exons code for protein sequences, they are important for the cell, yet constitute only ~2% of the sequences. Introns, in contrast, constitute 98% of the sequences but seem to have few crucial functions, except for enhancer sequences and developmental regulators in rare instances.
  Until Philip Sharp and Richard Roberts discovered introns within eukaryotic genes in 1977, it was believed that the coding sequence of all genes was always in one single stretch, bounded by a single long ORF. The discovery of introns was a profound surprise, which instantly brought up the questions of how, why and when the introns came into the eukaryotic genes.
  It soon became apparent that a typical eukaryotic gene was interrupted at many locations by introns, dividing the coding sequence into many short exons. Also surprising was that the introns were long, as long as hundreds of thousands of bases. These findings prompted the questions of why many introns occur within a gene (for example, ~312 introns occur in the human gene TTN), why they are long, and why exons are short.
  It was also discovered that the spliceosome machinery was large and complex with ~300 proteins and several SnRNA molecules. The questions extended to the origin of the spliceosome. Soon after the discovery of introns, it became apparent that the junctions between exons and introns on either side exhibited specific sequences that directed the spliceosome machinery to the exact base position for splicing. How and why these splice junction signals came into being was another important question.