How do sperm swim? How do they navigate? What is sperm made of? What does a World War Two codebreaker have to do with it all? The BBC untangles why we know so little about this mysterious cell.

With every heartbeat, a man can produce around 1,000 sperm – and during intercourse, more than 50 million of the intrepid swimmers set out to fertilise an egg. Only a few make it to the final destination, before a single sperm wins the race and penetrates the egg.

But much about this epic journey – and the microscopic explorers themselves – remains a mystery to science.

"How does a sperm swim? How does it find the egg? How does it fertilise the egg?" asks Sarah Martins da Silva, clinical reader of diabetes endocrinology and reproductive biology at the University of Dundee in the UK. Almost 350 years on from the discovery of sperm, many of these questions remain surprisingly open to debate.

Using newly developed methods, scientists are now following sperm on their migration – from their genesis in the testes all the way to the fertilisation of the egg in the female body. The results are leading to groundbreaking new discoveries, from how sperm really swim to the surprisingly big changes that occur to them when they reach the female body.

"Sperm – or spermatozoa – are 'very, very different' from all other cells on Earth," says Martins da Silva. "They don't handle energy in the same way. They don't have the same sort of cellular metabolism and mechanisms that we would expect to find in all other cells."

Due to the huge range of functions demanded of spermatozoa, they require more energy than other cells. Plus sperm need to be flexible, to be able to respond to environmental cues and varying energetic demands during ejaculation and the journey along the female tract, right up until fertilisation.

Sperm are also the only human cells which can survive outside the body, Martins da Silva adds. "For that reason, they are extraordinarily specialised." However, due to their size these tiny cells are very difficult to study, she says. "There's a lot we know about reproduction – but there's a huge amount that we don't understand."

One fundamental question that remained unanswered over almost 350 years of research: what exactly are sperm?

"The sperm is incredibly well-packaged," says Adam Watkins, associate professor in reproductive and developmental physiology at Nottingham University in the UK. "We typically thought of the sperm as a bag of DNA on a tail. But as we've started to realise, it's quite a complex cell – there's a lot of [other] genetic information in there."

The science of sperm began in 1677, when Dutch microbiologist Antoni van Leeuwenhoek looked through one of his 500 homemade microscopes and saw what he called "semen animals". He concluded, in 1683, that it wasn't the egg that contained the miniature and entire human, as previously believed, but that man comes "from an animalcule in the masculine seed". By 1685, he had decided that each spermatozoon contains an entire miniature person, complete with its own "living soul".

Almost 200 years later, in 1869, Johannes Friedrich Miescher, a Swiss physician and biologist, was studying human white blood cells collected from pus left on soiled hospital bandages when he discovered what he called "nuclein" inside the nuclei. The term "nuclein" was later changed to "nucleic acid" and eventually became "deoxyribonucleic acid" – or "DNA".

Aiming to further his studies of DNA, Miescher turned to sperm as his source. Salmon sperm, in particular, were "an excellent and more pleasant source of nuclear material" due to their particularly large nuclei. He worked in freezing temperatures, keeping laboratory windows open, in order to avoid deterioration of salmon sperm. In 1874, he identified a basic component of the sperm cell that he called "protamine". It was the first glimpse of the proteins that make up sperm cells. It took another 150 years, however, for scientists to identify the full protein contents of sperm.

Since then, our understanding of sperm has moved on leaps and bounds. But much still remains a mystery, says Watkins. As scientists have started to better understand early embryonic development, he adds, they are realising that sperm doesn't just pass the father's chromosomes on, but also epigenetic information, an extra layer of information that affects how and when the genes should be used. "It can really influence how the embryo develops and potentially the lifelong trajectory of the offspring that those sperm generate," says Watkins.

Sperm cells begin to form from puberty onwards, made in vessels within the testicles called seminiferous tubules.

"If you look inside the testes where the sperm are made, it starts as just a round cell that looks pretty much like anything else," says Watkins. "Then it undergoes this dramatic change where it becomes a sperm head with a tail. No other cell within the body changes its structure, its shape, in such a unique way."

It takes sperm

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