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Tragedy and triumph mixed on a Grand National day that elicited strongly contrasting emotions and raised questions about the race's future direction.

Just as those associated with winner Neptune Collonges had celebrated victory in the closest ever finish to the marathon Aintree contest, news emerged of two equine fatalities that cast a shadow over the event.

The head-to-head battle that saw the galloping grey deny Sunnyhillboy by a nose demonstrated what a thrilling spectacle the race can provide.

It was just the kind of drama that a sell-out crowd of 70,000, and a worldwide audience of 600 million, should savour.

But out on the track, Cheltenham Gold Cup winner Synchronised and According To Pete were put down after suffering falls.

You could sense the mood around the course changing, and racing's authorities were left to answer some searching questions about the nature of the contest afterwards.

The National has witnessed some extraordinary days, but few could have matched the events of the 165th running.

Top Irish jockey Ruby Walsh was ruled out of the race with a late injury for the second time in three years after a fall from Zarkandar in the Aintree Hurdle.

He was left to watch from the sidelines as his sister Katie sought to become the first woman to ride the National winner, on their father Ted's horse Seabass.

Walsh's great pal AP McCoy was aboard Synchronised, bidding to become the first horse since Golden Miller 78 years earlier to win steeplechasing's two biggest events in the same season.

But the 16-time champion jockey was unshipped before the start and the horse ran loose, before the pair were reunited and his mount was passed fit by a vet to race.

A delay of eight minutes and two false starts put nerves further on edge and then Synchronised came down at the course's fearsome Becher's Brook. He ran on without his rider but suffered a fractured leg five fences later.

According To Pete also fractured a leg having been brought down after jumping Becher's second time round, as just 15 of the 40 runners finished the race.

Walsh looked poised to make history in the four-and-a-half-mile contest but had to settle for third place, which was still the best ever finish by a female jockey.

Up ahead, Daryl Jacob brought Neptune Collonges home to become the first grey to win the race since Nicolaus Silver in 1961.

He got the verdict over Sunnyhillboy by the shortest possible distance, a nose, following a photo finish. The gallant 11-year-old was promptly retired on a winning note.

Jacob dedicated the win to former weighing-room colleague and housemate Kieran Kelly, who died in a fall in 2003.

"Kieran looked after me when I went to work for Dessie Hughes in Ireland and he virtually kicked me out of the house to tell me I had to go and ride in England so I wouldn't be here without him," said Jacob, 28.

It was a first National victory after 52 unsuccessful runners for champion trainer Paul Nicholls.

"This is a race we haven't had the best of luck in, but it's great to win. It's a race everybody knows and everyone watches," he said.

The Somerset trainer, whose horse Ornais was one of offered a robust defence of the National afterwards.

"When you are in competitive sport, whatever you do - motor racing, hockey - there is an element of risk. We take risks in everything we do in our lives, every single day," he said.

"All those people watch it because they enjoy watching the sport, but there are always risks. We all try and minimise them, and do what we can to help them.

"The worst thing you can do is to go too far. You make the fences smaller, they go faster and you get more fallers.

"We just have to get on with it, while making sure we do everything to make it as safe as possible. You can't make it 100% safe, but you must be sensible.

"Sport is risk. If people are going to continue to participate in sport, there is going to be both a human and animal risk.

"We have to live with that and get on with it. We have to grow up, basically. A lot of people have to grow up, and realise that it is life, and get on with it."

The margin between glory and despair was illustrated by the runner-up Sunnyhillboy's trainer Jonjo O'Neill.

One moment, he was inches away from winning the big race, and the next he learned his stable star Synchronised had died.

It bore haunting echoes of the day 33 years earlier when he rode Gold Cup winner Alverton, who died after a fall at Becher's Brook, and he was said to be inconsolable.

The National has been a national institution because of the drama and uncertainty as horses tackle 30 demanding fences.

Those who support it will point to the fact that the horses are bred to race, while opponents question whether fatalities are a price worth paying.

Aintree introduced several new safety measures following the deaths in the 2011 race, and will now be under pressure to go further.