Joshua Wong was 17 when he was first arrested for his political views. But by then he had been taking part in pro-democracy protests for more than three years. In 2011, aged 14, he founded a student activist group in Hong Kong to campaign against the government's introduction of a compulsory school curriculum that was favourable to the Communist Party of China: the new curriculum ignored events such as the 1989 Tiananmen Square protests in Beijing, in which several hundred students were shot and killed, and was critical of democracy.
In 2012, Wong organised large-scale demonstrations: a handful of students went on hunger strike and tens of thousands of people flooded the plazas outside Hong Kong's government headquarters. By 2014 – the year of his first arrest – Wong was a leading figure in the so-called Umbrella Revolution, a series of protests that swept across Hong Kong after China announced it would be screening candidates in the territory's own coming elections. Now 20, Wong is the secretary general of Demosisto, a pro-democracy political party he co-founded last year.
The problem is in the heart of the most mature democracies in the West – Joan Hoey, Economist Intelligence Unit
Wong is championed by democratic countries in the West. Time magazine nominated him for Person of the Year in 2014 and Fortune named him one of their “world's greatest leaders” in 2015. But as Wong – and others like him – fight for democracy, many countries that applaud his activism appear to be letting it slip. By more than one measure, democracy around the world is declining.
Trust in political institutions – including the electoral process itself - are at an all-time low. New converts to democracy in Europe and the Middle East are sliding back into authoritarian rule. And populist leaders who are expected to curb certain civil liberties are winning votes. Societies the world over are experiencing a strong backlash to a system of government that has largely been the hallmark of developed nations for generations.
“A lot of focus gets put on places like Russia, the Middle East or China,” says Joan Hoey at the Economist Intelligence Unit (EIU) in London. “But the problem is here, in the heart of the most mature democracies in the West.”
Hoey's stark assessment is shared by many others. Western states are worrying about the health of democracy for the first time since perhaps the end of World War Two, says Larry Diamond, a political sociologist at Stanford University in California and senior fellow at the Hoover Institution, a think tank that ‘promotes political and economic freedom’. “We’ve not seen anything like this in decades, and we don’t know where it’s heading,” he says. “We don’t know how serious it is.”
Diamond has been watching democracy around the world go through what he calls “a mild but protracted recession” for about a decade. Parts of the world new to the democratic system – such as former Soviet countries in Eastern Europe or states working through the aftermath of the Arab Spring – are slowly slipping back into authoritarianism.
We’ve not seen anything like this in decades, and we don’t know where it’s heading. We don’t know how serious it is
But last year, everything changed. Democracy is now in trouble in some of the most mature democracies in the world, he says. “We can now talk of a crisis.”
In fact, the decline of democracy has been measured. Every year since 2006, Hoey and her colleagues at the EIU have produced a report called the Democracy Index, which provides a comprehensive ranking of nearly every country in the world on a 10-point scale. It combines regional data and multiple surveys conducted in 167 countries to measure the quality of political processes, civil liberties, the functioning of government, public participation and political culture. Each country is then classed as a full democracy, flawed democracy, hybrid regime or authoritarian regime.
The results of
last year’s report are sobering. Overall, the global average score fell with 72 countries dropping in the ranking compared to 2015, and just 38 moving up. The number of “full democracies” dropped from 20 to 19, with the US now classed as “flawed”. According to the EIU's measure, around half the world's population (49.3%) live in a democracy of some kind. But only 4.5% of people live in a “full democracy” - half as many as in 2015.
And the EIU's measure is not the only one that suggests a rapid, fundamental shift in global politics. Andrew Reynolds, a political scientist at the University of North Carolina and founder of the Electoral Integrity Project, which assesses the quality of democracies around the world, has argued that the US state of
North Carolina should no longer be considered a democracy after it brought in voting restrictions that reportedly disenfranchised black voters.
So, what's going on? What’s behind the erosion of a political system that’s guided the world’s most developed economies for decades?
A common explanation is that the world is still reacting to the global financial crisis and the austerity policies that followed. This had a major corrosive effect on democracy, changing the way people viewed their political leaders. According to this view, the effect will be short-term – when economies start to pick up again, politics will return to normal. But what we're seeing is not a temporary blip, says Hoey.
Take the US. Its relegation to “flawed democracy” in the EIU’s ratings is not because of the 2016 presidential election. “The US has been teetering on the brink for many years,” says Hoey. “Donald Trump is a beneficiary of a deep-seated and long-standing problem.”
The level of public trust in democratic institutions in the US has been plummeting for decades. According to a survey carried out in 2015 by the Pew Research Center, a nonpartisan organisation in Washington DC that investigates demographic trends, only
19% of people trust the government to do the right thing “always or most of the time”. In 1958, when the American National Election Study asked the same question, 73% of people did.
Some may argue that this is because governments no longer feel like they are “of the people, by the people, for the people”, as Abraham Lincoln put it in his Gettysburg Address. Over the last half century, the business of governing has arguably become more technocratic, with positions of power populated by larger numbers of professional politicians and policy wonks. Many long-established political parties once had closer ties with specific groups of people. Left-wing or social democratic parties in particular were set up to represent the will of the working class. Those ties have stretched to breaking point, however.
More generally, old divisions between left and right that once gave voters clear alternatives have fallen, especially since the 1990s and the end of the Cold War. Parties that represented two competing visions of how society should be run throughout the 20th Century have suffered a body blow, says Hoey. As parties on both sides moved to the centre, the gulf between political elites and the electorate opened up even more. “Politics is no longer about the big questions and big issues,” says Hoey. “It has become soulless.”
Cue populists like Donald Trump and Nigel Farage, the former leader of UK party Ukip. Such politicians have been able to win support by talking about issues that established parties have been unwilling to address candidly. Ukip wields no hard political power – its only elected member of Parliament defected last week – but its outspoken views on immigration and criticism of EU technocrats shaped the Brexit debate. Similarly, Trump also crafted his campaign around immigration and a pledge to “drain the swamp” of political elites that no longer shared the values of millions of voters.
Many are suddenly talking about the need to defend democracy. 'But defend democracy against what? Against the people?'
The resulting political shocks were a wake-up call, says Hoey. But in failing to talk about the things that mattered to people, the mainstream parties had it coming. People want their voices heard and when they had an opportunity to make a difference with a direct vote – one that promised to make a bigger difference than the usual box-ticking every four years - they grabbed it. “The chickens have come home to roost,” she says.
As a result of the populist backlash, political elites – which includes many in the media – are suddenly talking about the need to defend democracy. “But defend democracy against what? Against the people?” asks Hoey. By getting the public involved in the biggest political debate in decades, Brexit was phenomenal, she says. “People who hadn’t voted for years came out.”
Yet many still identify the populist backlash itself as the problem, rather than an expression of a deeper issue. Brexit and Trump voters are stigmatised for being bigots – “deplorables” – or for being misled by misinformation or lying politicians. But to dismiss millions of people like that will get us nowhere, says Hoey.
“Our political parties have run away from talking about the issues that matter to people,” she says. “If you're not asking the really big questions about what kind of society you want to live in, what’s left?” If people care about something, it needs to be discussed – no matter how difficult a topic.
“You need to have clashes of opinion,” she argues. “If you want to revise democracy that’s the only way to do it. There are no other fixes.”
For Hoey, Brexit and the election of Trump are electoral shocks that could be good for democracy in the long-run. “All these years, nobody’s really cared about democracy,” she says. “Suddenly everyone’s talking about it and that’s great.” But Diamond sees a darker side. “Many deep thinkers about politics, from Plato to the authors of the US constitution, have worried about the vulnerability of pure democracy to the tyranny of the majority.”
Future Now asked 50 experts – scientists, technologists, business leaders and entrepreneurs – to name what they saw as the most important issues of the 21st Century.
The reason that many countries have representative democracies – in which people elect representatives to make decisions on their behalf – or other structures, such as second chambers of government, is that the will of the people needs to be balanced with things like equality and civil liberty. Some states have constitutions that set out their citizens' incontrovertible rights explicitly. Most have independent judicial systems. “You need brakes, like in a car,” says Diamond. “If all you have is the accelerator pedal it’s not a very safe vehicle.”
Many deep thinkers about politics have worried about the vulnerability of pure democracy to the tyranny of the majority
The danger inherent in the democratic process is that a leader can be elected who removes those brakes. When people feel threatened, either physically – by terrorism, say – or economically, they tend to be more receptive to authoritarian populist appeals and more willing to give up certain freedoms.
Trump has support for banning immigration from certain Muslim-majority countries, for example. And last year, the UK government was able to pass the most sweeping internet-surveillance legislation of any democracy. “In the US and most of Western Europe, the checks and balances are very likely to be strong enough to prevent severe damage to democratic freedoms and constitutional safeguards,” says Diamond. “But 'very likely' is not 'certain'.”
Diamond is struck by how quickly democratic processes and institutions are being dismantled in European countries like Hungary and Poland – states that have long been absorbed into the European Union. “Maybe we are going to have some shocking lessons about the durability of democracy,” he says.
Diamond agrees with Hoey about the underlying causes of the populist surge across the West. “When people are saying they can't stomach any more immigration, when they don't know if they're going to be able to retire or what kind of jobs their kids are going to get, the political elite needs to listen and adapt or things are going to unravel,” he says.
But simply talking about these issues may not be enough. To compete with more authoritarian rivals, Diamond thinks mainstream politicians will need to concede ground, stepping back from liberal social and economic policies – on equality, immigration or global trade – that have been advanced in recent years. For example, Geert Wilders' nationalist party in the Netherlands did worse than expected in this month’s elections. This was because the Dutch prime minister saw what was happening and made some significant policy adjustments, says Diamond.
Despite being on the back foot, many people believe democracy is the best system of government humans have come up with, an end point to political evolution. In non-democratic countries around the world – in parts of Asia, in sub-Saharan Africa –
survey data shows that people want it. As China has become richer and its economy more modern, you can see a growing aspiration for democracy from the middle classes, says Hoey. “It's human nature to want to be free.”
The political elite needs to listen and adapt or things are going to unravel
Which is why people like Joshua Wong devote themselves to fighting for it. Feelings are strong on both sides, however. When Wong travelled to Taiwan in January, he was met by around 200 pro-China protestors at the airport. One broke through police lines and tried to punch him. Wong ended up being placed under police protection for his visit. Is democracy really the only morally legitimate system for choosing a society's leaders?
“I don’t see any stable authoritarian states out there,” says Diamond. He believes governments in places like China, Russia and Iran will eventually collapse. ”The only well-functioning authoritarian regime in the world is Singapore and I'm not sure even that is going to last,” he says. “In any case, you can’t build a theory on a city state of just a few million people.”
Not everyone thinks things are so clear cut, however. Daniel Bell at Tsinghua University in Beijing argues that a lot of Western ideas about democracy verge on dogma. A Canadian political scientist trained in the UK, Bell has spent many years living and working in China. “In the West we tend to divide the world into good democratic regimes that set the path for all the others, and bad authoritarian regimes that are on the wrong side of history,” he says.
Bell points out that non-democratic states can take many forms. There are family-run dictatorships like in North Korea, military dictatorships like in Egypt, monarchies like in Saudi Arabia. Each is quite different. And some, like China's meritocratic system – in which government officials are not elected by the public, but appointed and promoted according to their competence and performance – should not be dismissed outright. “To put them all in the same camp is ridiculous,” says Bell. “It’s not a good way of trying to understand what’s going on in China.”
The Communist Party of China has 88 million members. Its membership is managed by the Department of Organisation, which is essentially a huge human resources department. To be a member of the party, candidates must pass a set of examinations. Government officials are thus selected from across the country and from various sectors of society according to merit. Promotion from low-ranking official to the very top of government is then – in principle – simply a matter of performance.
One obvious issue is a lack of transparency in how merit is measured. At the lower levels of government, the system is becoming more open to public scrutiny. Some Chinese cities are now experimenting with putting budgets online and allowing people to comment on the budgets, which lets citizens see how their local officials are performing. But how the party selects its top-tier leaders is not generally known, says Bell. “If they were a bit more open, it would help to legitimise the system abroad.”
If [China] say they’re going to do something by 2030, we can be pretty sure they’re going to do it
The biggest challenge to Chinese politics is corruption. A democratic system can live with corruption because corrupt leaders can be voted out of power, at least in theory. But in a meritocratic system, corruption is an existential threat. If political leaders are seen to be corrupt, they cannot claim superior merit and thus lose the one quality that justifies their position. Because of this, China needs more mechanisms to keep its politicians accountable. Chinese officials have studied the British civil service to learn how to deal with corruption, for example. “Elections are a safety valve that isn't available in China,” says Bell. “But they know this. It's why they're having the longest and most systematic anti-corruption drive in recent history.”
There are obvious flaws in China's system, says Bell. But he also ticks off several advantages. Political officials at the top all have substantial experience at running a country – “unlike in the US with the current president”. The government is also not subject to the electoral cycle and can focus on its policies. “If they say they’re going to do something by 2030, we can be pretty sure they’re going to do it,” he says.
This has allowed China to pull millions out of poverty in just a few decades, build a vast amount of new infrastructure in the
biggest construction drive the world has ever seen, and begin to tackle its substantial urban pollution and greenhouse emissions. Officials used to be judged mainly on how well they did at reducing poverty, says Bell. Now they are expected to make environmental improvements too.
The West has tried to export democracy not only at the point of a gun, but also by imposing legislation
Bell says lots of surveys show that the Chinese system has strong support within the country at most levels of society, where the government is viewed as providing a form of guardianship. He agrees with Hoey that as China gets richer and its middle class grows, more people will want to have a say in how the country is run. But that need not necessarily mean a call for democracy. Instead, perhaps more people will sign up to join the ruling party. Everybody now has equal rights to take the examinations that put you on the road to becoming a public official, he says. “There are different ways of participating in politics.”
Whatever happens, democracy is much more likely to flourish when it is homegrown. The attempts in the last few decades to export democracy around the world have proved to be an absolute disaster, says Hoey. “The whole idea is wrong in principle because democracy is not ours to dispense,” she says. “It has to come from the people to have any meaning. It needs to have roots deep in the values and culture of the country.”
Yet the West has tried to export democracy not only at the point of a gun – such as in the many military interventions in the Middle East – but also by imposing legislation. The EU pushes its Western values and body of laws on new members, for example. This can be quite intrusive, says Hoey. As a result, rather than being seen as a universal human aspiration, democracy can sometimes come across as a specifically Western product – and rejected as such.
With the political climate around the world shifting and many countries adopting a more nationalist outlook, the US and Western Europe have abandoned most of their ambitions for regime change around the world. But looking inwards may be no bad thing. “If the West wants to promote democracy then they should do it by example,” says Hoey.
Many thousands of years ago, not a single human being lived in the Americas.
This only changed during the last Ice Age. It was a time when most of North America was covered with a thick sheet of ice, which made the Americas difficult to inhabit.
But at some point during this time, adventurous humans started their journey into a new world.
They probably came on foot from Siberia across the Bering Land Bridge, which existed between Alaska and Eurasia from the end of the last Ice Age until about 10,000 years ago. The area is now submerged by water.
There is still debate about when these first Americans actually arrived and where they came from. But we are now getting closer to uncovering the original narrative, and finding out who these first Americans really were.
During the peak of the last Ice Age about 20,000 years ago, a journey from Asia into the Americas would not have been particularly desirable. North America was covered in icy permafrost and tall glaciers. But, paradoxically, the presence of so much ice meant that the journey was, in a way, easier than it would be today.
The abundance of ice meant that sea levels were much lower than they are now, and a stretch of land emerged between Siberia and Alaska. Humans
and animals could simply walk from Asia to North America. The land bridge was called Beringia.
People were using the woody shrubs from the land bridge to ignite bones on the landscape
At some point around this time – known as the Last Glacial Maximum – groups of hunter-gatherers moved east from what is now Siberia to set up camp there.
"The first people who arrived in Beringia were probably small, highly mobile groups evolving in a large landscape, probably depending on the availability of seasonal resources," says
Lauriane Bourgeon of the University of Montreal, Canada.
These people did well to seek refuge there. Central Beringia was a much more desirable environment than the icy lands they had left behind. The climate
was a bit damper. Vegetation, in the form of woody shrubs, would have given them access to wood that they could burn to keep warm.
Beringia was also an ideal environment for large grazing mammals, giving early hunter-gathers something to hunt, says
Scott Elias of Royal Holloway, University London in the UK, who reconstructs past climates.
"Our hypothesis is that people were using the woody shrubs from the land bridge to ignite bones on the landscape. The bones of big animals contain lots of fatty deposits of marrow, and they will burn."
When humans got to Beringia, they would have had little choice but to set up camp there. The vast Laurentide and Cordilleran ice sheets further east cut them off from North America.
This standstill helped these isolated groups of people to become genetically distinct from those they had left behind
It is now becoming clear that they made Beringia their home, staying put for several thousand years. This idea is called the Beringian Standstill Hypothesis. This standstill helped these isolated groups of people to become genetically distinct from those they had left behind, according to a
2007 study.
This long standstill therefore meant that the people who arrived in the Americas – when the ice finally retreated and allowed entry – were genetically different to the individuals who had left Siberia thousands of years earlier. "Arguably one of the most important parts of the process is what happened in Beringia. That's when they differentiated from Asians and started becoming Native Americans," says
Connie Mulligan of the University of Florida in Gainesville, US, who took part in this early analysis.
Since then, other genetic insights have further supported the standstill hypothesis.
Elias and colleagues even propose that people stayed in Beringia for as long as 10,000 years.
When the ice finally started to retreat, groups of people then travelled to different pockets of the Americas.
There has long been debate over whether these early settlers arrived from several migrations from different areas, or just one.
There's been no turnover or change in the population group as some people had previously hypothesised
Over 20 years ago,
Mulligan proposed that there was just one migration from Beringia into the "New World". She came to this conclusion by analysing the genetic variation in the DNA of modern-day Native Americans and comparing it with the variation in Asia. The same rare pattern appeared in all the Native Americans she studied, but very rarely appeared in modern-day Asians. This meant Native Americans likely arose from a single population of people who had lived in Beringia, isolated for many years.
In 2015,
a study using more advanced genetic techniques came to a similar conclusion.
Rasmus Nielsen of the University of California, Berkeley, US, and colleagues found that the "vast majority" of Native Americans must have originated from just one colonisation event.
"There's been no turnover or change in the population group as some people had previously hypothesised," says Nielsen. In fact, about 80% of Native Americans today are
direct descendants of the Clovis people, who lived across North America about 13,000 years ago. This discovery came from a 2014 genetic study of a one-year-old Clovis boy who died about 12,700 years ago.
But we now know there must have been staggered migrations from Beringia.
That is because there are small groups of people in the Amazonian region of South America – such as the Suruí and Karitiana – with additional mysterious "arctic gene flow", unrelated to the Clovis boy.
Another 2015 study therefore proposed there was more than one "founding population of the Americas".
The indigenous populations of the Americas, the team found, have distant genetic links in common with people of Australia, Papua New Guinea and the Andaman Islands.
People came into Beringia over different times during the standstill
This means, says
Pontus Skoglund of Harvard University in Boston, Massachusetts, that people came into Beringia over different times during the "standstill" and went on to populate different parts of the Americas. Those early dispersals are still reflected by differences in the genomes of people living today.
"It wasn't simply a single homogenous founding population. There must have been some type of patchwork of people, and maybe there were multiple pulses," says Skoglund.
In other words, the Beringian inhabitants did not all arrive or leave at the same time.
This makes sense when you consider that Beringia was not a narrow land bridge with ocean on either side. "It was a huge region about twice the size of Texas," says Elias. The people living there would have had no idea that it was a land bridge at all. "There were no sign posts saying they were leaving Siberia."
This makes it highly likely that there were different groups of Beringian inhabitants that never met.
A study
published in February 2017 strengthens this idea further. After examining the shapes of 800- to 500-year-old skulls from Mexico, researchers found they were so distinct, the people the skulls belonged to must have remained genetically isolated for at least 20,000 years.
To understand who the first Americans really were, we have to consider when they arrived. While the exact timing is hard to pin down. Nielsen's work gives some insight. By sequencing the genomes of people from the Americas, Siberia, and Oceania, he and colleagues could understand when these populations diverged. The team concludes that the ancestors of the first Americans came to Beringia at some point between 23,000 years and 13,000 years ago.
We found cut marks on bones from horse, caribou and wapiti so we know that humans were relying on those species
We now have archaeological evidence to suggest that the people who left Siberia – and then Beringia – did so even earlier than the 23,000-year-limit proposed by Nielsen and colleagues. In January 2017, Lauriane Bourgeon and her team found evidence of people living in a cave system in the northern Yukon Territory of western Canada, called the Bluefish Caves,
that dates to as early as 24,000 years ago. It was previously believed that people had only arrived in this area 10,000 years later.
"They reached Beringia as early as 24,000 years ago, and they remained genetically and geographically isolated until about 16-15,000 years ago, before dispersing south of the ice sheets that covered most of North America during this period," says Bourgeon.
The caves "were only used on brief occasions for hunting activities", she says. "We found cut marks on bones from horse, caribou and wapiti, so we know that humans were relying on those species."
This work provides further evidence that people were in the Beringia area at this early date. But it does not reveal the exact dates these people first ventured further south.
For that, we can turn to archaeological evidence. For decades, stone tools left by the Clovis people
have been found throughout North America. Some date to as early as 13,000 years ago. This might suggest that humans moved south very late. But in recent years evidence has begun to emerge that questions this idea.
Most preserved remains are stone tools and sometimes bones of animals
"We really know little about them, because most preserved remains are stone tools and sometimes bones of animals, thus technology and diet," explains
Tom Dillehay of Vanderbilt University in Tennessee, US, who is studying these people. "Monte Verde in south-central Chile, where I am at present, has several organic remains – animal hide, meat, plant remains that reveal a wider diet, wood technology – but these types of sites are rare to find."
Another conundrum remains. Ice sheets still covered North America 18,500 years ago, making journeying south difficult. How did people arrive in southern Chile so early?
A leading idea had been that an ice-free corridor opened up, which allowed humans to travel south. However, the latest evidence suggests this corridor only opened
about 12,600 years ago, long after these early Chileans arrived.
Elias also points out how difficult this journey would have been. "Even if there was a small gap in between these enormous ice sheets, the environment left in that gap would have been so horrible, with mud, ice, meltwater and slush. It would not have been a habitable place for people or the animals they would have wanted to follow," he says.
These early people could have travelled by boat
There is an alternative. These early people could have travelled by boat, taking a route along the Pacific coast. There is no archaeological evidence to support this idea, but that is not entirely unexpected: wooden boats are rarely preserved in the archaeological record.
There are still many unanswered questions, but Mulligan says that studying how and when early hunter-gatherers spread across the Americas can help us to understand the process of migration itself. That is, how population sizes change and which genetic traits persist.
In many ways, the peopling of America presents scientists with a golden opportunity to study these processes. There have been multiple migrations both into and out of other regions of the world – Africa, Europe and Asia, for instance. But the people who moved into the Americas were on a one-way journey. "We know the original inhabitants came from Asia into the New World with no other people there, and no major back migrations, so it's the simplest model you can conceive of."
That it was a one-way journey, coupled with the increased interest in studying the genetics of these ancient people, means we should soon understand even more about who these first Americans really were, and exactly when they arrived.
In August 2016, researchers estimated a five-metre-long female
Greenland shark had lived for 392 years, making it the longest-lived vertebrate. The mammalian lifespan record belongs to a bowhead whale, thought to have reached the grand old age of 211.
Perhaps it is because humans have become so dominant in other respects that we are fascinated by species that outlive us. For biologists, examples of extreme longevity raise fundamental questions about why organisms age and die. And given that they do, why can individuals of some species live for hundreds of years while others get months, weeks or even just days?
Humans are relatively long-lived. Some researchers hope that gaining greater knowledge of what drives longevity in the animal kingdom offers the chance, not only to understand those species better, but our own too. Others go further, believing it is the key to longer, healthier human lives.
The discovery of Ming's extraordinary age in 2013 led to
immediate speculation that the secret to its long life lay in its very low oxygen consumption.
Indeed, one of the most deeply-entrenched ideas about animal lifespan is that it is closely linked to metabolic rate – or the speed of chemical reactions that break food down into energy and produce compounds needed by cells. The notion that animals undergo cumulative damage and die sooner when they work harder like machines run at full capacity probably dates back to the Industrial Revolution.
Why can individuals of some species live for hundreds of years while others get months, weeks or even just days?
In the early 20th Century, German physiologist Max Rubner compared rates of energy metabolism and lifespans in guinea pigs, cats, dogs, cows, horses and humans. He found that larger animals had lower metabolic rates per gram of tissue and that they lived longer, leading him to conclude that using up energy faster shortened life.
American biologist Raymond Pearl developed the idea further following his research on the effects of starvation, temperature change and heredity on the lifespans of fruit flies and cantaloupe melon seedlings. "In general the duration of life varies inversely as the rate of energy expenditure during life," he wrote in his 1928 book
The Rate of Living.
In 1954 Denham Harman, at the University of California, Berkeley, provided a mechanism to support what became known as the rate of living theory. He proposed ageing to be the result of an accumulation of damage caused to cells by free radicals. Generated during metabolism, these are highly reactive molecules that can damage cellular machinery by stealing electrons.
However, while it is true that larger species of mammals have slower metabolic rates and live longer, the rate of living theory has largely been abandoned. For one thing, researchers have pointed out many birds and bats live much longer than they should for their metabolic rates.
Marsupials have shorter lifespans than placental mammals despite having lower metabolic rates.
John Speakman of the University of Aberdeen in the UK is among those who have highlighted that, just because animals with slower metabolic rates have longer lifespans, does not mean the former causes the latter.
"All the evidence that has been used to support the rate of living theory has a fundamental flaw in it," says Speakman. "That is, it comes from studies which compared animals with different body sizes."
For mammals, once you take out the influence of body size, it's those with higher metabolic rates that live longer
In 2005, Speakman used a clever statistical trick to remove the influence of body mass from the equation,
in a study of data for 239 mammalian species and 164 species of birds. For each animal with a higher-than-expected metabolic rate for its body size, he examined whether it had a correspondingly lower-than-expected life span for its body size, and vice versa. "For both mammals and birds, once body mass was removed, the relationship between metabolic rate and life span was zero," says Speakman.
However this calculation, like previous work supporting the rate of living theory, used the resting metabolic rates of animals, when they are neither digesting food nor regulating body temperature. Researchers have traditionally used these rates simply because more data is available for animals in this state. However, many animals spend only a minority of their time at a resting rate of metabolism, and the proportion of time different species spend at it varies widely.
"It turns out there is a relationship, but it's the opposite of what you predict from the rate of living theory," says Speakman. "For mammals, once you take out the influence of body size, it's those with higher metabolic rates that live longer." The results for birds did not reach statistical significance.
In fact, the idea that the more oxygen an animal consumes, the greater the production of free radicals that cause damage, and therefore the swifter the ageing, is now outdated. That is thanks to more detailed studies of mitochondria, the parts of cells that generate energy.
When mitochondria break down chemicals within food, protons are pushed across their inner membranes, creating a difference in electrical potential across them. When the protons are released back across the membrane, this potential difference is used to create adenosine triphosphate (ATP), a molecule which stores energy.
It was originally thought that free radical production is high when the electrical difference across the mitochondria membrane was high – meaning that the higher the rate of metabolism, the greater the production of highly reactive molecules that cause cellular damage and ageing.
Smaller animals have more predators, and have to grow faster, as well as reproduce sooner
In fact this model fails to take account of the presence of "uncoupling proteins" in the mitochondria inner membrane. With functions including heat generation, these uncoupling proteins trigger the flow of protons across the membrane to reduce the potential difference across it when it is high.
"The traditional idea that, as you increase your metabolism, a fixed percentage of the oxygen you are consuming will go off to produce free radicals, is fundamentally at odds with what we know about the way mitochondria work," says Speakman. "If anything, we would expect that as metabolism goes up and uncoupling goes up… free radical damage would go down."
Across these four groups, Magalhães found that 63% of the variation in lifespan was down to body mass. For mammals only, it was 66%. Bats are something of an outlier in that they live much longer than they should for their size, so he re-worked the calculation without them, and this time he found body mass explained 76% of mammal lifespan variation. The association for birds was 70% and for reptiles it was 59%. There was no correlation for amphibians.
Magalhães and others who have studied the impact of size on how long animals live say it comes down to combined evolutionary and ecological factors.
"Body size is a determinant of ecological opportunities," says Magalhães. "Smaller animals have more predators, and have to grow faster, as well as reproduce sooner, if they want to pass on their genes. Larger animals, like elephants and whales, are less likely to be eaten by predators, and lack the evolutionary pressure to mature and reproduce at an early age."
The island opossums lived on average four-and-a-half months, or 23%, longer than their mainland cousins
If body size affects lifespan via likelihood of being eaten, it follows that different populations of the same species could live for longer or shorter periods in different environments.
Steven Austad, a journalist-turned-lion-tamer-turned-biologist, set out to test this idea in a study of adult female opossums in the late 1980s. He caught and attached radio collars to 34 on Sapelo Island, Georgia, US, and to another 37 on the mainland near Aitken, South Carolina, US. The second of these populations is hunted by wild dogs and bobcats (
Lynx rufus), while the population on the island is not. The island opossums are under less pressure from predators generally, and are genetically isolated.
Austad found the island opossums lived on average four-and-a-half months, or 23%, longer than their mainland cousins. They also had significantly smaller litters, began reproducing a little later and were able to reproduce for longer. Tests showed that collagen in tail tendon fibres aged more quickly in the mainland opossums.
There are other factors that at first glance might seem to have an impact on species lifespan, but in fact turn out to be just a function of body size and ecological opportunities. Brain size, for example, has been shown to correlate with maximum species lifespans,
especially in primates, as has eyeball size. "If you have anything that changes with body size, it will look as if it is related to lifespan, simply because there is a relationship between body size and lifespan," says Speakman.
While there is a prevailing scientific consensus around the importance of body size on lifespans via likelihood of being killed by other animals, this still leaves vital questions unanswered.
"It depends on the level at which you ask the question," says Speakman. "The evolutionary explanation is to do with extrinsic mortality risk. The question then is what are the actual mechanisms that protect the body?"
A mutation in a gene called daf-2 is known to allow nematode worms to live doubled yet still healthy lifespans
In his hunt for answers to this question, Austad turned, in research published in 2010, to a group of long-lived animals he called
Methusaleh's Zoo, after the biblical patriarch said to have lived for 969 years. Austad argued that the low-temperature environments of longevity record holders such as Ming the clam, Greenland sharks and bowhead whales are no coincidence.
"Most animals that live an exceptionally long time have a low body temperature, or live in a low-temperature environment," he says. Austad points out that key bodily processes such as reactive oxygen species production, DNA repair and gene transcription are slower in the cold.
Being especially interested in processes that could inform human lifespan extension, Austad also paid special attention to naked mole rats and little brown bats, two mammals that outlive humans relative to body mass. He concluded that the accumulation of damage to cells as a result of the production of free radicals does play a role in ageing, but one that is relatively minor in many cases, and that varies in importance between species.
The development of quick, cheap DNA sequencing technologies in recent years has offered scientists important clues about the roles of genes in regulating longevity in a variety of species. For example, a mutation in a gene called
daf-2 is known to allow nematode worms to
live doubled yet still healthy lifespans. Dwarf mice with mutated versions of genes that undermine production of growth hormone, the hormone prolactin and thyroid-stimulating hormone,
live about 40% longer than control animals.
What explains the surprisingly low rates of cancers in large, long-lived animals like elephants and whales?
In 2015, he went on to lead a group that
sequenced the genome of the bowhead whale, revealing species-specific mutations in genes linked to DNA damage response, the regulation of cell cycles and the control of cancer.
"We don't know for a fact that these are the proteins involved in species differences in ageing, but these studies offer clues we can take forward and test further," says Magalhães. He is currently involved in an international collaboration that is sequencing the capuchin monkey, which can live past the age of 50, despite its relatively small size.
Magalhães and others gathering this growing database of the genetic determinants of longevity are seeing a pattern in the enhanced DNA repair capabilities of long-lived animals. For instance, sequencing has solved a biological mystery that has puzzled scientists since the 1970s; what explains the surprisingly low rates of cancers in large, long-lived animals like elephants and whales?
In 2015, a team led by Joshua Schiffman, of the University of Utah,
calculated that fewer than 5% of captive elephants die from cancers, compared to a cancer mortality rate of 11-25% in humans. When they looked at data from sequencing studies, they found the African elephant has 40 copies of the gene that encodes p53 – a protein that plays a key anti-cancer role, by either preventing cells with damaged DNA from dividing until repairs have been performed, or triggering them to commit suicide. Asian elephants have 30 to 40 copies. Both humans and the rock hyrax, elephants' closest living relative, have just two copies of the gene.
Being long-lived is part of what makes us human, yet we don't understand why we have that capacity
"My hypothesis is that it's not DNA repair capacity per se that is different, rather it's the way cells respond to DNA damage," says Magalhães. "The same amount of DNA damage is going to kill an elephant cell or stop it proliferating, but not necessarily a mouse cell."
"It would make little evolutionary sense for short-lived animals to waste valuable energy defending themselves against diseases that take many years to develop," says Austad. "It would be like putting a $1,000 face on a cheap watch."
Scientists using comparative biology to understand ageing now have access to the genomes of dozens of mammals. As this increases to hundreds, they will be better able to identify genetic clues to the drivers of longevity.
"Being long-lived is part of what makes us human, yet we don't understand why we have that capacity," says Magalhães. "Sequencing more species will help us find out, and to answer many other fascinating questions."