The technique has therefore been heavily influenced by the development of high-throughput sequencing technologies.[2][3] Massively parallel signature sequencing (MPSS) was an early example based on generating 16–20 bp sequences via a complex series of hybridisations,[38][note 1] and was used in 2004 to validate the expression of ten thousand genes in Arabidopsis thaliana.[39] The earliest RNA-Seq work was published in 2006 with one hundred thousand transcripts sequenced using 454 technology.[40] This was sufficient coverage to quantify relative transcript abundance.