Horsemeat was unknowingly sold by supermarkets across the UK after a gang of three men secretly mixed it with beef and then falsified documents, a court has heard.
The specific chain of events that led horsemeat to enter the food chain, sending shockwaves through the British food industry, has been publicly outlined for the first time in the trial of a businessman said to have played a "key role" in the scandal.
Andronicos Sideras, 55, is alleged to have mixed horsemeat with beef and then forged paperwork to sell it on as "100 per cent beef" to manufacturers who made products for well-known retailers including Tesco, Co-op and Asda.
The contamination was first identified in Ireland before tests by the Food Standards Agency found horsemeat in products on sale in the UK.
At the height of the furore, thousands of items were pulled from the shelves as supermarkets were forced to apologise amid serious concerns about the food supply chain.
Jonathan Polnay, prosecuting, told jurors at Inner London Crown Court: "This case, stripped to its essentials, is actually very straightforward.
"It is about lying to people. It is about deceiving people to make money. Or to be more precise - for people to make more money.
"Like most if not all offences of dishonesty, it was motivated by greed."
Mr Sideras, from London, was arrested in 2013 alongside Ulrik Nielsen, 57, the owner of Danish company FlexiFoods and his "right-hand man", Alex Ostler-Beech, 44, from Hull.
All three were charged with conspiracy to defraud and the latter two admitted their part in the 2012 scam, jurors heard.
Mr Sideras, who denies the charge, co-owned Dinos & Sons, a meat company and sausage manufacturer in Tottenham, north London.
The court heard that FlexiFoods bought horsemeat and beef from suppliers across Europe and had it delivered to Dinos & Sons, which allegedly charged up to four times more than usual storage costs to mix the meats into "horsebeef".
At the time, beef sold for around three euros per kilogram at wholesale prices, while horsemeat was cheaper at two euros per kilogram.
Mr Polnay said: "Whilst at Dinos - and this is the heart of our case - the horsemeat and the beef would be mixed together into a single load.
"Dinos would create false paperwork and labels to make it look like all the meat being supplied was beef."
Meanwhile, FlexiFoods, which has an office in Hull, would sell the mixed meat to Martin McAdam, owner of meat trading company McAdam Food Products, based in Ireland, the jury was told.
Mr Polnay said it was not clear whether Mr McAdam was aware he was buying "horsebeef".
He added: "Mr McAdam had the contacts to sell the meat on to large meat production companies that make products for a vast range of well-known companies in this country and in fact all across Europe.
"It goes without saying that they would be told they were buying beef. They would not be interested in horsemeat."
FlexiFoods would then arrange for the mixed meat to be delivered directly from Dinos & Sons to the "large production companies".
Mr Polnay said that for the fraud to work it "needed someone" to physically mix the meat.
He added: "It needed someone to fix the documents, to make them look genuine.
"We say that key role was taken by this defendant - Andronicos Sideras."
Mr Polnay said a trail of falsified invoices, ledgers and email conversations between Mr Sideras and FlexiFoods were "almost direct evidence of the conspiracy".
"They are meticulous in their records," he added. "You can see how they account for their fraud."
He told jurors: "There is no dispute that this fraud was going on - the sole question you are going to need to decide is whether Mr Sideras was involved in it."
A joint probe carried out by City of London Police, the Food Standards Agency and the CPS has taken just over three years to complete.
In January 2015, abattoir owner, Peter Boddy, 65, from Todmorden, West Yorks, became the first person to be convicted over the scandal. He pleaded guilty to failing to comply with food traceability regulations over 17 horse carcasses sold for meat.
The trial of Mr Sideras, expected to last four weeks, continues.