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The Jambos made the ideal start when they went ahead through Lawrence Shankland’s early penalty, but the visitors equalised with a spot-kick of their own shortly afterwards.

Hearts enjoyed the bulk of the pressure thereafter and Shankland saw a goal ruled out for offside just before Andrija Zivkovic’s superb 75th-minute strike against the run of the play handed the Greeks the initiative.

The cinch Premiership side now face the formidable task of needing a victory at the hostile Toumba Stadium in Thessaloniki next Thursday if they are to prolong their European run into the group stage.

Hearts made four changes to the team that started Sunday’s 4-0 Viaplay Cup victory over Partick Thistle as Odel Offiah, Toby Sibbick, Calem Nieuwenhof and Alex Lowry were replaced by Nathaniel Atkinson, Stephen Kingsley, Cammy Devlin and Liam Boyce.

Backed by another packed Tynecastle crowd, the hosts started on the front foot and there were loud shouts for a penalty in the sixth minute when Kenneth Vargas fell under a challenge from Rafa Soares as he burst into the PAOK box.

Latvian referee Andris Treimanis initially was not convinced but after VAR intervened and following a lengthy check of the pitchside monitor, he pointed to the spot. Three minutes after the foul was committed, Shankland, having kept his composure amid the tension, slotted his kick into the net.

The hosts’ celebrations were short-lived, however. Just two minutes after going ahead, they let the Greeks back into it when Kye Rowles chopped down Serbia forward Zivkovic in the box.

It was a clear penalty and captain Stefan Schwab duly tucked his kick beyond Zander Clark, much to the delight of the partisan 300-strong band of travelling fans.

Hearts almost regained the lead in the 31st minute when Devlin’s cross into the box was glanced towards goal by the head of Shankland but goalkeeper Dominik Kotarski did superbly to claw it out. Vargas and Shankland both had attempts to force the rebound in before Boyce, who looked certain to score from six yards out on the follow-up, saw his shot blocked by Soares.

The Jambos threatened again two minutes into the second half when Kingsley flashed an angled half-volley just beyond the far post after PAOK failed to deal with Alex Cochrane’s free-kick into the box.

The Edinburgh side – who lost Barrie McKay to injury just after the hour – thought they had gone ahead in the 66th minute when Shankland curled a superb finish into the top corner after being played in by Boyce just inside the box but after a VAR check, and following lengthy celebrations from the hosts, the scorer was ruled to be offside.

It proved a pivotal moment as PAOK got themselves ahead in the 75th minute when Zivkovic fired home a ferocious low strike from just outside the box after a corner kick was worked into his path, sparking wild celebrations among the visiting players and supporters.