Category: Domestication

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The apples of our eye

Apples have been another recurring obsession on the blog, maybe because they make the conservation problem unusually tangible, and immediate. Pretty much everyone thinks they know what an apple is, but the diversity hidden behind that word is enormous, and not much visible in supermarkets. A lot of it may have disappeared, but you can pretty much guarantee a slew of heritage apple stories in the press come autumn (and so it begins), so I guess people care. Here’s a selection of posts about apples spanning 15 years. There’s a lot more out there in the archive for you to explore.

Let’s start at the beginning, or as close to it as the genetics can get us. In 2016’s The Adam and Eve of apples?, I discussed the wild apple forests of southeastern Kazakhstan, where all of the world’s apple varieties can be traced back to an ancestral population of Malus sieversii — spread, if you believe the more colourful tellings, by bears eating the sweetest fruit and pooping the seeds around the forest. As usual, though, the genetics turned out to be more interesting than the headline. The cultivated apple isn’t a simple straight line from one mother tree; it’s the product of a long history of movement, hybridization and exchange between wild and domesticated populations all along the old trade routes out of Central Asia.

But those forests are in danger. Back in 2009, in Wild fruit relatives threatened in Central Asia, I looked at how the wild relatives of apples and other crops were vanishing along with their habitat. Among them was Malus niedzwetzkyana, the remarkable red-fleshed apple of Kyrgyzstan. These aren’t just obscure wild trees: they represented genetic diversity that has already made its way into cultivated apples, and might still have something to offer breeders. That red-fleshed apple came back a year later in Not the world’s first red apple. A new red-fleshed variety was being breathlessly presented in the press as a novelty — not GM, just 15 years of patient crossing and 20,000 rejected seedlings — but nature had got there first. M. niedzwetzkyana has been doing red flesh for millennia.

The cultivated stock has its own story of loss. In 2009’s Saving an apple a day, I looked at the loss of American apple varieties, and at the people trying to rescue what remained. Thousands of named varieties had once been grown in the United States; only a fraction survived in orchards, collections and the memories of the people who knew what they tasted like. It was a nice illustration of a familiar conservation paradox: you can lose a lot of diversity without even noticing, because the supermarket keeps selling you apples.

However, that same year, a study added a wrinkle to the “thousands lost” narrative. In Lost apples found, Jeremy pushed back a little on the standard version of the story — the one where 6,800 of 7,100 named 19th-century American apple varieties are simply gone. A team from the University of Arizona and USDA had gone looking for ghosts among old apple trees on historic farmsteads in Arizona, Utah and New Mexico. Only 34 of the resulting genotypes matched a known variety. The other 110, spread across 160 trees, matched nothing on record. Some of those are probably unnamed seedlings that never had a name to lose in the first place. But some, surely, are the supposedly lost varieties everyone had given up on. Not extinct, just lurking.

Seven years on, the USDA apple folks came back with another version of the answer. In As American as apples, Gayle Volk summarized her and Adam Henk’s trawl through historic nursery catalogues, which turned up 891 named apple cultivars grown in the US before 1908. They went looking for the 150 that seemed to matter most. The payoff: 90% are still findable today, safely tucked away in genebanks, private collections or specialist nurseries. Only 51 high-priority cultivars remained unaccounted for in any collection. For now.

The lesson isn’t that genetic erosion doesn’t happen. It plainly does, and 30 fruit nurseries vanishing a year is not nothing. No, the lesson is that “extinct” is much harder to prove than “endangered,” and pomologists with their equivalent of a swab kit keep turning out to be more patient, and luckier, than we have any right to hope.

Conserving what’s left, of course, depends on people as much as on collections and gene sequencers. In 2014, I offered up Irish eyes smiling about apple conservation. The post was prompted by Michael Hennerty’s beautiful book on the Irish apple, but the real story was the people who had nurtured the 68 varieties documented in the book over the years. Conservation often depends on a motley collection of people keeping things alive long enough for some fancy institution to discover that they matter. A few years later, Jeremy went into much more detail on Irish apples on his podcast.

Institutions, though, can be just as precarious as the work of any orchard-keeper or amateur gardener. Across the Irish Sea, the story of the UK’s apple collection showed just how fragile even a major national genebank can be. The National Fruit Collection at Brogdale in Kent, with its hundreds of apple varieties, came under threat when its funding was thrown into doubt, prompting a succession of stories about sales, new management and relocations. Oh, and royalty. Anyway, the collection is fine, but the saga was a reminder that conservation is not just about keeping things alive. You have to keep them alive with enough institutional resilience that they can survive the next crisis.

Taken together, these posts highlight a sometimes overlooked aspect of safeguarding agrobiodiversity. Conservation is not just putting varieties safely into collections, nor, on the other hand, is it just protecting wild populations from destruction. Apples show that the two are connected — or should be. These are, in the jargon, complementary conservation strategies. The diversity in a genebank has a history outside it, and the diversity out in the wild and in farms may only have a future in a genebank. We really do need both, it’s not just mom and apple pie rhetoric.

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Brainfood: Archaeology edition

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Crops made, and remade

Two new studies of very different crops – banana and chrysanthemum, of all things – end up telling surprisingly similar, deliciously complicated stories. For these crops, domestication was not once and done, so to speak. They were both repeatedly remade as people moved them through landscapes containing new wild diversity.

In the banana study, the authors suggest that a partly domesticated Musa acuminata lineage from New Guinea was carried westwards through Southeast Asia, encountering and hybridizing again and again with different local wild bananas along the way. Each encounter added new genetic material to an already changing crop, helping produce the genomic mosaics found in mainland Southeast Asian bananas today.

The chrysanthemum study reveals a strikingly parallel history: cultivated plants originating in China were introduced to Japan, where they encountered local wild populations and acquired new genetic diversity, before later movements to Europe and further breeding reshaped the crop again.

The papers also show that an essentially similar process played out somewhat differently in the two crops. In banana, repeated hybridization appears to have been important in the building of the crop itself, as domesticated or partly domesticated plants became the starting material for successive encounters with wild Musa. In chrysanthemum, genomic analysis reveals a more complex network of relationships among multiple wild and cultivated groups, with C. indicum among the important ancestral contributors. In this genetic cauldron, hybridization and introgression repeatedly diversified an established cultivated genepool, contributing to traits such as flower form, colour and plant architecture.

Taken together, these papers challenge the familiar “funnel,” or bottleneck, image of domestication: a one-way downward slide from diverse wild relative to genetically narrow crop. Instead, they point to a more stop-start, two-way, non-linear, geographically contingent process, in which (semi-)cultivated plants continue to encounter, absorb and be reshaped by wild diversity.

In both banana and chrysanthemum — and probably many other cases — wild relatives have been more active, continuous participants in creating the diversity of the crops we know today than we sometimes give them credit for. Or I have given them credit for at any rate.

That argues for treating the wild genepool not simply as a reservoir from which to fish out useful genes one at a time. The historical evidence suggests that crops have benefited in the past from repeatedly absorbing larger chunks of wild genetic diversity, allowing selection to reconstruct useful combinations. Might it be worth trying to make that happen again? Is anyone out there doing pre-breeding explicitly with an eye to the past?

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The fruits of migration

As chance would have it, following my recent speculative foray into the possible connection between language and crop diversity, an essay by the great Andrea Wulf 1 has just appeared that tells the story of how a remarkable German naturalist used breadfruit and linguistics to work out Polynesian migration patterns long before DNA or archaeology could confirm them.

George Forster sailed with his father aboard the HMS Resolution on Cook’s second voyage (1772-1775) and encountered breadfruit across the South Pacific. He noticed that the trees were seedless and could only be propagated by root cuttings. And also that there didn’t seem to be wild breadfruit trees anywhere in the islands. That meant every tree he saw, and there were many of them, had been deliberately planted by people.

Digging through the university library back in Göttingen, Forster later found that wild, seeded breadfruit with little flesh do exist, but in Southeast Asia. If the Pacific islands only had the improved, seedless types, he reasoned, they must have been carried there by people. That was proof both of sophisticated Polynesian plant management and of a migration route running from Southeast Asia eastward across the Pacific.

Forster combined this with careful attention to language. He deduced that most Pacific languages he encountered were dialects of a single root language, with small, systematic sound shifts. He and his father also charted how the presence of Malay loanwords decreased the farther east an island was located, which suggested that islands closer to Southeast Asia — the home of Malay — were settled earlier.

Where his fellow naturalist Joseph Banks saw the breadfruit purely as an economic resource (later lobbying to transplant it to the West Indies to feed enslaved plantation workers, hence HMS Bounty and all that), Forster saw it as historical and ethnographic evidence. That’s a strikingly different, more humanistic and — to me at least — more appealing way of reading a plant.

Wulf then jumps forward to modern archaeology and genetics, which broadly confirm Forster’s reconstruction. There was an initial human population in the region of Papua New Guinea dating back 45,000 years. This was followed by a second, Austronesian-speaking wave migrating out of Taiwan and thereabouts around 3,500 years ago. They travelled by the stars in “outrigger or double-hulled canoes loaded with hogs, chickens, root vegetables, coconuts, bananas, and breadfruit seedlings.” And they mixed with the descendants of the earlier wave through Melanesia. Their own descendants eventually reached Fiji, Samoa, Tonga, and, much later, Tahiti, Hawaii, Easter Island and New Zealand.

The essay closes by tying this to Tongan historian Epeli Hau’ofa’s idea of “Oceania” — the Pacific not as scattered, isolated islands, lost in a vast ocean, but as one interconnected “sea of islands.” Exactly what Forster intuited two centuries before it became conventional wisdom.

We’ve blogged a lot about breadfruit over the years. Check out the archive. And in particular listen to the recently-retired Dr Diane Ragone describe her 40-year journey to understand this remarkable tree.

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A Babel of agrobiodiversity

“The rise and fall of language diversity through the Holocene” by Blasi, Hamilton, Gray & Bowern, published in Science last month, does something I really love. It complicates a standard narrative.

The received story of language diversity is that it has been declining steadily for centuries, largely driven by colonialism and globalization. But direct evidence is scarce and recent: writing only reliably takes us back about 6,000 years. So the team combined ethnographic data from 171 hunter-gatherer and fisher societies, paleodemographic estimates of global population, and statistical and social-computational modelling to reconstruct plausible trajectories of global language diversity over the 12,000 years of the Holocene, which began after the last major ice age, triggering the birth of agriculture.

At the beginning of this period, they estimate there were roughly 4,500–6,200 languages. That’s actually fewer than the roughly 7,500 languages spoken today. Language diversity then increased through most of the Holocene, driven by the population growth engendered by agriculture, reaching a peak between about 1,000 and 3,000 years ago, when the models suggest that tens of thousands of languages may have existed. That’s an order of magnitude more than today.

Then came a major collapse. The authors argue that the decline in linguistic diversity began with the expansion of large states and empires, long before European colonialism. Expanding populations and their languages, cultures and institutions displaced smaller ones. Today’s languages are therefore “survivors of a massive and highly selective historical bottleneck.” They persist because the populations speaking them expanded. Extinction, the authors argue, has shaped linguistic and cultural diversity far more — and for longer — than we have appreciated.

Now, there are obvious caveats. The authors are modelling linguistic diversity, not measuring it. Their estimates depend on heroic assumptions about early ethnolinguistic group sizes, global population and the relationship between population and language number. Regional histories could have been very different, and the model can’t reconstruct individual episodes of extinction caused by war, disease or environmental change.

But the paper provides a provocative template for thinking about diversity more generally. It suggests that population growth can initially generate diversity; but expanding societies can subsequently erase it through homogenization.

And that makes me wonder about agricultural biodiversity. You saw that coming, I’m sure.

I’m not suggesting that crop diversity followed the same pattern as language diversity through the Holocene. We simply don’t have the evidence to say that, though it’s an intriguing thought. But could we borrow the methodology? Could we reconstruct plausible trajectories of crop or agricultural-system diversity from fragmentary archaeological, ethnobotanical, genetic and demographic evidence?

Rather than assuming that agricultural diversity was vast and stable in the distant past, happy in its Vavilovian centres, and has been declining recently, perhaps we might end up complicating the familiar genetic erosion narrative too. That would be worthwhile. And fun.