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with the much smaller Chinese vari- ety (), which has some natural immunity to the Asian . And after decades of work, he is within reach of his goal, a tall American with enough Chinese traits to keep it healthy. Other researchers are trying to attack the blight with viruses or are creating that are genetically modified (GM) to resist the fungus, and could be the first GM forest trees released in the wild in the . Progress with all three approaches is raising hopes that chest- nuts will soon start to flourish again in the for- ests of the American east. “We’re starting to pull the tree back from the brink of extinction,” says Hebard. The work is also offering lessons that could COURTESY OF THE FOREST HISTORY SOCIETY, DURHAM, NORTH CAROLINA SOCIETY, HISTORY OF THE FOREST COURTESY help to save other trees, such as elm and ash, which face similar threats in America and abroad. FROM GIANTS TO STUMPS Once known as the sequoia of the east, the American chestnut was one of the tallest trees in the forest, and dominated a range of 800,000 square kilometres, from to (see ‘Felled by a fungus’). It made up 25% of the forest, and its annual was a major source of food for both animals and humans. The decay-resistant was also used to make telephone poles, roofs, fence posts and parts of railway lines. The first warning signs came in 1904, when rust-coloured cankers developed on at in New York. Zoo Before the fall: American chestnuts in the Great Smokey Mountains of North Carolina in 1910. forester Hermann Merkel took a sample across the street to the New York Botanical , where mycologist William Murrill soon iden- tified the as . The blight probably hitched a ride on nursery imports of Japanese chestnuts beginning in 1876. Spread- The chestnut ing through rain and air, fungal spores infected trees through bark wounds and breaks. Can- kers developed, quickly encircling a branch or trunk and cutting off the supply of water and resurrection nutrients from the soil. Within 50 years, the blight had laid waste to nearly the entire popu- Once king of eastern forests, the American chestnut was lation of some 4 billion trees. Other hardwoods, mainly , eventually wiped out by blight. Now it is poised to rise again. filled the void but they do not produce a con- sistent crop of nuts every year. “You had a really BY HELEN THOMPSON species of eastern . With long, dominant species that the wildlife depended straight trunks and bushy crowns, it carpeted on which was then replaced with a species that hey’re hard to breed and easy to kill,” the forest floor each autumn with prickly then didn’t produce as much,” says Douglass says pathologist Fred Hebard as brown nuts. But the arrival of chestnut blight Jacobs, a forest ecologist at Purdue University “T he attacks a 2-metre-tall chestnut tree ( parasitica) from wiped out in West Lafayette, Indiana. Reports from the in southwest . Hebard bores a hole in almost all the stately trees, leaving only a few, period suggest that squirrel populations ini- the bark and squeezes a mash of orange fun- isolated stands. Since then, a faithful fan club tially collapsed, and that five moth species gus into the wood. The tree is a of the of scientists and citizens has sought to tame dependent on chestnuts went extinct (D. A. Chinese and American chestnut species, and the blight. Orwig J. Biogeogr. 29, 1471–1474; 2002). Hebard hopes that it has enough resistance As chief scientist of the American Chest- As the chestnut succumbed to the blight, genes to keep the fungus — called chestnut nut Foundation (ACF), a group of chestnut scientists raced to find solutions. They started blight — at bay. If so, the hybrid could help to enthusiasts and scientists, Hebard has bred breeding hybrids of American and Asian resurrect a long-gone icon. thousands of hybrids at the organization’s chestnuts, which have evolved alongside the Until a century ago, the American chestnut research farm in Meadowview, Virginia. He blight. But the attempts produced no trees that (Castanea dentata) was the cornerstone tree crosses descendants of the original American were sufficiently resistant yet still had enough

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American traits to make them a popular Chuck Maynard have planted some 600 trans- also concerned about the threat of diseases replacement. Asian chestnuts are shorter and genic trees for field trials of disease resistance. already attacking other trees, such as root less sturdy than their American counterparts. A transgenic variety with a gene for the rot mould ( cinnamomi) and Over the years, chestnut lovers have tried enzyme oxalate oxidase, which disarms the fun- invasive insects, including ambrosia beetles a range of remedies, including , gus, has already shown resistance in the field. and gall wasps. Hybridization and transgenic sulphur fumes and radiation. They even con- Maynard, Powell and their colleagues are programmes are beginning to target some of ducted religious ceremonies, assuming that the collaborating with a non-profit organization these threats. sins of the people had brought on the blight. called the Forest Health Initiative in Research Lessons learned from the battle to restore But all these strategies failed. Triangle Park, North Carolina, to develop a the chestnut could help save other threatened GM version of an American chestnut with trees. The American elm () RESTORATION CHESTNUT strong resistance based on genes from Asian and its European counterpart (Ulmus laevis) In 1983, a group of plant scientists joined together to form the ACF with the goal of “We’re starting to pull the American chestnut creating a resistant tree. The foundation has since grown to 6,000 volunteer members, rang- back from the brink of extinction.” ing from retired physicists to farmers, and it maintains 486 regional breeding and chestnuts. Because such ‘cisgenic’ trees contain have been hit hard by Dutch elm disease, and 120,000 experimental trees. only chestnut genes, researchers hope that the ash and hemlock are succumbing to invasive At Meadowview Research Farm, where trees won’t provoke strong public objection. insects. In the United Kingdom, the beloved Hebard is based, a refrigerator holds prized The Forest Health Initiative is testing the cis- horse chestnut () is in pollen from American chestnuts and between genic chestnuts now and aims to obtain federal danger of being wiped out by a combination of 600 and 700 strains of the fungus. The valley approval to conduct larger plantings. “Chest- a bacterium and a moth. Researchers are devel- and hillsides are lined with rows of hybrid nut may be the first case of a genetically engi- oping hybrid and transgenic programmes to chestnuts as well as unadulterated American, neered tree that’s planted out. If that happens it defend against many of those threats. Chinese and Japanese chestnuts (Castanea cre- can probably pave the way for other trees,” says nata). Some have plastic bags covering their Jacobs. Along the way, the transgenic work will NEVER-ENDING BATTLE flowers to prevent unwanted natural pollina- help researchers to determine which resistance “It’s the pest of the month club. We’ve lost chest- tion from ruining the experiments. “It’s a chas- genes are the most useful. nut and lost elm. Now, it’s almost a new species tity belt for chestnuts,” jokes Hebard. “You bag Other researchers are enlisting viruses that or pest is being identified and a new tree or it if you want to know who the daddy is.” attack the chestnut fungus. Such viruses spread forest is being threatened almost on a monthly Nearby are rows of chestnuts that stand most easily among closely related fungi and are basis,” says Carlton Owen, a forester and wildlife about a metre tall, the most promising hybrid effective at controlling the blight in . biologist who chairs the Forest Health Initiative’s yet to emerge from nearly 30 years of breed- But American fungal strains are more diverse, governing board. Owen says that the results of ing experiments. This ‘restoration chestnut’ is so the virus cannot spread as effectively. To get the chestnut work will help researchers to pro- 94% American and 6% Chinese and seems to around that problem, virologist Don Nuss at tect other species by showing which strategies show strong resistance to the blight. But the the University of Maryland in College Park has work best in different situations and by testing Virginia trees may not thrive in other loca- developed a transgenic fungus that is designed public acceptance of GM trees. tions, so the researchers are working to adapt to spread the virus more easily. Other research- Once researchers have a resistant chestnut, the restoration chestnut to other climates. At ers are conducting field trials to test whether the question is where to plant it. Forest eco- the same time, they are breeding new hybrids the engineered fungus can compete with — systems have transformed in the past century, at other experimental farms to preserve a and replace — wild-type fungal strains. If so, and reintroducing the chestnut could upset healthy degree of genetic diversity. the viruses could be used in targeted ways to the new ecological balance. “You can’t assume More than 1,100 kilometres to the north, protect American chestnuts. that a forest with chestnut is better than a for- researchers are experimenting with chestnuts Most researchers agree that restoring the est without. You can’t roll the clock back,” says that contain genes thought to provide resistance, American chestnut will require a combination Steve , chief scientist at the Environ- which were taken from Chinese chestnuts as of fungal viruses and resistant trees, whether mental Defense Fund in Boston. well as such as wheat, peppers and grapes. hybrid, GM or a mixture of both. And the “Reintroduction is going to be kind of a At the State University of New York in Syracuse, trees will need to defend themselves against gradual thing,” says Jacobs. In 2009, the ACF plant pathologist Bill Powell and forest biologist more than just the blight. Researchers are began planting restoration chestnuts on US Forest Service land in Virginia, and North Carolina. In April 2012, it also started FELLED BY A FUNGUS planting hybrid chestnuts and other hardwood Before an Asian fungus wiped them out, chestnuts dominated saplings at a former mining site. eastern forests of the United States. For chestnut lovers, other signs of hope stand alongside a path at the New York Botanical Garden. In a corner of the garden, New York: Fungal blight a transgenic chestnut and a restoration hybrid rst spotted both reach about a metre high. Although in 1904 their have shrivelled a bit, the scrappy Nashville saplings have managed to survive one of the Washington DC warmest summers on record. And their bark is smooth, with no sign of the cankers that Natural range claimed their ancestors. ■ of the American Atlanta Chestnut Helen Thompson recently completed an internship at Nature in Washington DC.

4 OCTOBER 2012 | VOL 490 | NATURE | 23 © 2012 Macmillan Publishers Limited. All rights reserved