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Thursday, May 16, 2019

Forest

From Wikipedia, the free encyclopedia

Forest on Mount Dajt, Albania
 
A forest is a large area dominated by trees. Hundreds of more precise definitions of forest are used throughout the world, incorporating factors such as tree density, tree height, land use, legal standing and ecological function. According to the widely used Food and Agriculture Organization definition, forests covered 4 billion hectares (9.9×109 acres) (15 million square miles) or approximately 30 percent of the world's land area in 2006.

Forests are the dominant terrestrial ecosystem of Earth, and are distributed around the globe. Forests account for 75% of the gross primary production of the Earth's biosphere, and contain 80% of the Earth's plant biomass. Net primary production is estimated at 21.9 gigatonnes carbon per year for tropical forests, 8.1 for temperate forests, and 2.6 for boreal forests.

Forests at different latitudes and elevations form distinctly different ecozones: boreal forests around the poles, tropical forests around the Equator, and temperate forests at the middle latitudes. Higher elevation areas tend to support forests similar to those at higher latitudes, and amount of precipitation also affects forest composition.

Human society and forests influence each other in both positive and negative ways. Forests provide ecosystem services to humans and serve as tourist attractions. Forests can also affect people's health. Human activities, including harvesting forest resources, can negatively affect forest ecosystems.

Definition

Forest in the Scottish Highlands
 
Although forest is a term of common parlance, there is no universally recognised precise definition, with more than 800 definitions of forest used around the world. Although a forest is usually defined by the presence of trees, under many definitions an area completely lacking trees may still be considered a forest if it grew trees in the past, will grow trees in the future, or was legally designated as a forest regardless of vegetation type.

There are three broad categories of forest definitions in use: administrative, land use, and land cover. Administrative definitions are based primarily upon the legal designations of land, and commonly bear little relationship to the vegetation growing on the land: land that is legally designated as a forest is defined as a forest even if no trees are growing on it. Land use definitions are based upon the primary purpose that the land serves. For example, a forest may be defined as any land that is used primarily for production of timber. Under such a land use definition, cleared roads or infrastructure within an area used for forestry, or areas within the region that have been cleared by harvesting, disease or fire are still considered forests even if they contain no trees. Land cover definitions define forests based upon the type and density of vegetation growing on the land. Such definitions typically define a forest as an area growing trees above some threshold. These thresholds are typically the number of trees per area (density), the area of ground under the tree canopy (canopy cover) or the section of land that is occupied by the cross-section of tree trunks (basal area). Under such land cover definitions, an area of land can only be known as forest if it is growing trees. Areas that fail to meet the land cover definition may be still included under while immature trees are establishing if they are expected to meet the definition at maturity.

Under land use definitions, there is considerable variation on where the cutoff points are between a forest, woodland, and savanna. Under some definitions, forests require very high levels of tree canopy cover, from 60% to 100%, excluding savannas and woodlands in which trees have a lower canopy cover. Other definitions consider savannas to be a type of forest, and include all areas with tree canopies over 10%.

Some areas covered in trees are legally defined as agricultural areas, e.g. Norway spruce plantations in Austrian forest law when the trees are being grown as Christmas trees and below a certain height.

Etymology

Since the 13th century, the Niepołomice Forest in Poland has had special use and protection. In this view from space, different coloration can indicate different functions.
 
The word forest comes from Middle English, from Old French forest (also forès) "forest, vast expanse covered by trees"; first introduced in English as the word for wild land set aside for hunting without the necessity in definition for the existence of trees. Possibly a borrowing (probably via Frankish or Old High German) of the Medieval Latin word foresta "open wood", foresta was first used by Carolingian scribes in the Capitularies of Charlemagne to refer specifically to the king's royal hunting grounds. The term was not endemic to Romance languages (e.g. native words for "forest" in the Romance languages evolved out of the Latin word silva "forest, wood" (English sylvan); cf. Italian, Spanish, Portuguese selva; Romanian silvă; Old French selve); and cognates in Romance languages, such as Italian foresta, Spanish and Portuguese floresta, etc. are all ultimately borrowings of the French word. 

A forest near Vinitsa, North Macedonia
 
The exact origin of Medieval Latin foresta is obscure. Some authorities claim the word derives from the Late Latin phrase forestam silvam, meaning "the outer wood"; others claim the term is a latinisation of the Frankish word *forhist "forest, wooded country", assimilated to forestam silvam (a common practice among Frankish scribes). Frankish *forhist is attested by Old High German forst "forest", Middle Low German vorst "forest", Old English fyrhþ "forest, woodland, game preserve, hunting ground" (English frith), and Old Norse fýri "coniferous forest", all of which derive from Proto-Germanic *furhísa-, *furhíþija- "a fir-wood, coniferous forest", from Proto-Indo-European *perkwu- "a coniferous or mountain forest, wooded height".

Uses of the word "forest" in English to denote any uninhabited area of non-enclosure are now considered archaic. The word was introduced by the Norman rulers of England as a legal term (appearing in Latin texts like the Magna Carta) denoting an uncultivated area legally set aside for hunting by feudal nobility.

Tywi Forest, Wales
 
These hunting forests were not necessarily wooded much, if at all. However, as hunting forests did often include considerable areas of woodland, the word "forest" eventually came to mean wooded land more generally. By the start of the 14th century, the word appeared in English texts, indicating all three senses: the most common one, the legal term and the archaic usage. Other terms used to mean "an area with a high density of trees" are wood, woodland, wold, weald, holt, frith and firth. Unlike forest, these are all derived from Old English and were not borrowed from another language. Some classifications now reserve the term woodland for an area with more open space between trees and distinguish among woodlands, open forests, and closed forests based on crown cover.

Evolution

The first known forests on Earth arose in the Late Devonian (approximately 380 million years ago), with the evolution of Archaeopteris. Archaeopteris was a plant that was both tree-like and fern-like, growing to 10 metres (33 ft) in height. Archaeopteris quickly spread throughout the world, from the equator to subpolar latitudes. Archaeopteris formed the first forest by being the first known species to cast shade due to its fronds and forming soil from its roots. Archaeopteris was deciduous, dropping its fronds onto the forest floor. The shade, soil, and forest duff from the dropped fronds created the first forest. The shed organic matter altered the freshwater environment, slowing it down and providing food. This promoted freshwater fish.

Ecology

Forests account for 75% of the gross primary productivity of the Earth's biosphere, and contain 80% of the Earth's plant biomass. Forest ecosystems can be found in all regions capable of sustaining tree growth, at altitudes up to the tree line, except where natural fire frequency or other disturbance is too high, or where the environment has been altered by human activity. 

The latitudes 10° north and south of the equator are mostly covered in tropical rainforest, and the latitudes between 53°N and 67°N have boreal forest. As a general rule, forests dominated by angiosperms (broadleaf forests) are more species-rich than those dominated by gymnosperms (conifer, montane, or needleleaf forests), although exceptions exist.

Forests sometimes contain many tree species within a small area (as in tropical rain and temperate deciduous forests), or relatively few species over large areas (e.g., taiga and arid montane coniferous forests). Forests are often home to many animal and plant species, and biomass per unit area is high compared to other vegetation communities. Much of this biomass occurs below ground in the root systems and as partially decomposed plant detritus. The woody component of a forest contains lignin, which is relatively slow to decompose compared with other organic materials such as cellulose or carbohydrate.

Components

Even, dense old-growth stand of beech trees (Fagus sylvatica) prepared to be regenerated by their saplings in the understory, in the Brussels part of the Sonian Forest.
 
A forest consists of many components that can be broadly divided into two categories that are biotic (living) and abiotic (non-living) components. The living parts include trees, shrubs, vines, grasses and other herbaceous (non-woody) plants, mosses, algae, fungi, insects, mammals, birds, reptiles, amphibians, and microorganisms living on the plants and animals and in the soil.

Layers

 
Spiny forest at Ifaty, Madagascar, featuring various Adansonia (baobab) species, Alluaudia procera (Madagascar ocotillo) and other vegetation
 
A forest is made up of many layers. Starting from the ground level and moving up, the main layers of all forest types are the forest floor, the understory and the canopy. The emergent layer exists in tropical rainforests. Each layer has a different set of plants and animals depending upon the availability of sunlight, moisture and food.
  • Forest floor contains decomposing leaves, animal droppings, and dead trees. Decay on the forest floor forms new soil and provides nutrients to the plants. The forest floor supports ferns, grasses, mushroom and tree seedlings.
  • Understory is made up of bushes, shrubs, and young trees that are adapted to living in the shades of the canopy.
  • Canopy is formed by the mass of intertwined branches, twigs and leaves of the mature trees. The crowns of the dominant trees receive most of the sunlight. This is the most productive part of the trees where maximum food is produced. The canopy forms a shady, protective "umbrella" over the rest of the forest.
  • Emergent layer exists in the tropical rain forest and is composed of a few scattered trees that tower over the canopy.

Types

A dry sclerophyll forest in Sydney, which is dominated by eucalyptus trees.
 
Forests can be classified in different ways and to different degrees of specificity. One such way is in terms of the biome in which they exist, combined with leaf longevity of the dominant species (whether they are evergreen or deciduous). Another distinction is whether the forests are composed predominantly of broadleaf trees, coniferous (needle-leaved) trees, or mixed.
The number of trees in the world, according to a 2015 estimate, is 3 trillion, of which 1.4 trillion are in the tropics or sub-tropics, 0.6 trillion in the temperate zones, and 0.7 trillion in the coniferous boreal forests. The estimate is about eight times higher than previous estimates, and is based on tree densities measured on over 400,000 plots. It remains subject to a wide margin of error, not least because the samples are mainly from Europe and North America.

Forests can also be classified according to the amount of human alteration. Old-growth forest contains mainly natural patterns of biodiversity in established seral patterns, and they contain mainly species native to the region and habitat. In contrast, secondary forest is regrowing forest following timber harvest and may contain species originally from other regions or habitats.

Different global forest classification systems have been proposed, but none has gained universal acceptance. UNEP-WCMC's forest category classification system is a simplification of other more complex systems (e.g. UNESCO's forest and woodland 'subformations'). This system divides the world's forests into 26 major types, which reflect climatic zones as well as the principal types of trees. These 26 major types can be reclassified into 6 broader categories: temperate needleleaf; temperate broadleaf and mixed; tropical moist; tropical dry; sparse trees and parkland; and forest plantations. Each category is described as a separate section below.

Temperate needleleaf

Temperate needleleaf forests mostly occupy the higher latitude regions of the Northern Hemisphere, as well as high altitude zones and some warm temperate areas, especially on nutrient-poor or otherwise unfavourable soils. These forests are composed entirely, or nearly so, of coniferous species (Coniferophyta). In the Northern Hemisphere pines Pinus, spruces Picea, larches Larix, firs Abies, Douglas firs Pseudotsuga and hemlocks Tsuga, make up the canopy, but other taxa are also important. In the Southern Hemisphere, most coniferous trees (members of the Araucariaceae and Podocarpaceae) occur in mixtures with broadleaf species, and are classed as broadleaf and mixed forests.

Temperate broadleaf and mixed

Broadleaf forest in Bhutan
 
Temperate broadleaf and mixed forests include a substantial component of trees in the Anthophyta. They are generally characteristic of the warmer temperate latitudes, but extend to cool temperate ones, particularly in the southern hemisphere. They include such forest types as the mixed deciduous forests of the United States and their counterparts in China and Japan, the broadleaf evergreen rainforests of Japan, Chile and Tasmania, the sclerophyllous forests of Australia, central Chile, the Mediterranean and California, and the southern beech Nothofagus forests of Chile and New Zealand.

Tropical moist

There are many different types of tropical moist forests, with lowland evergreen broad leaf tropical rainforests, for example várzea and igapó forests and the terra firma forests of the Amazon Basin; the peat swamp forests, dipterocarp forests of Southeast Asia; and the high forests of the Congo Basin. Seasonal tropical forests, perhaps the best description for the colloquial term "jungle", typically range from the rainforest zone 10 degrees north or south of the equator, to the Tropic of Cancer and Tropic of Capricorn. Forests located on mountains are also included in this category, divided largely into upper and lower montane formations on the basis of the variation of physiognomy corresponding to changes in altitude.

Tropical dry

Tropical dry forests are characteristic of areas in the tropics affected by seasonal drought. The seasonality of rainfall is usually reflected in the deciduousness of the forest canopy, with most trees being leafless for several months of the year. However, under some conditions, e.g. less fertile soils or less predictable drought regimes, the proportion of evergreen species increases and the forests are characterised as "sclerophyllous". Thorn forest, a dense forest of low stature with a high frequency of thorny or spiny species, is found where drought is prolonged, and especially where grazing animals are plentiful. On very poor soils, and especially where fire or herbivory are recurrent phenomena, savannas develop.

Sparse trees and parkland

Taiga forest near Saranpaul in the northeast Ural Mountains, Khanty–Mansia, Russia. Trees include Picea obovata (dominant on right bank), Larix sibirica, Pinus sibirica, and Betula pendula.
 
Sparse trees and savanna are forests with lower canopy cover of trees. They occur principally in areas of transition from forested to non-forested landscapes. The two major zones in which these ecosystems occur are in the boreal region and in the seasonally dry tropics. At high latitudes, north of the main zone of boreal forest, growing conditions are not adequate to maintain a continuous closed forest cover, so tree cover is both sparse and discontinuous. This vegetation is variously called open taiga, open lichen woodland, and forest tundra. A savanna is a mixed woodland grassland ecosystem characterized by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach the ground to support an unbroken herbaceous layer consisting primarily of grasses. Savannas maintain an open canopy despite a high tree density.

Forest plantations

Forest plantations are generally intended for the production of timber and pulpwood. Commonly mono-specific and/or composed of introduced tree species, these ecosystems are not generally important as habitat for native biodiversity. However, they can be managed in ways that enhance their biodiversity protection functions and they can provide ecosystem services such as maintaining nutrient capital, protecting watersheds and soil structure, and storing carbon.

Societal significance

Redwood tree in northern California redwood forest, where many redwood trees are managed for preservation and longevity, rather than being harvested for wood production
 
Forests provide a diversity of ecosystem services including
Some researchers state that forests do not only provide benefits, but can in certain cases also incur costs to humans. Forests may impose an economic burden, diminish the enjoyment of natural areas, reduce the food producing capacity of grazing land  and cultivated land, reduce biodiversity reduce available water for humans and wildlife, harbour dangerous or destructive wildlife, and act as reservoirs of human and livestock disease.

The management of forests is often referred to as forestry. Forest management has changed considerably over the last few centuries, with rapid changes from the 1980s onwards culminating in a practice now referred to as sustainable forest management. Forest ecologists concentrate on forest patterns and processes, usually with the aim of elucidating cause-and-effect relationships. Foresters who practice sustainable forest management focus on the integration of ecological, social, and economic values, often in consultation with local communities and other stakeholders.

Priest River winds through mountains with a checkerboard design of trees to its east
Priest River winding through Whitetail Butte with lots of forestry to the east—these lot patterns have existed since the mid-19th century. The white patches reflect areas with younger, smaller trees, where winter snow cover shows up brightly to the astronauts. Dark green-brown squares are parcels
 
Humans have generally decreased the amount of forest worldwide. Anthropogenic factors that can affect forests include logging, urban sprawl, human-caused forest fires, acid rain, invasive species, and the slash and burn practices of swidden agriculture or shifting cultivation. The loss and re-growth of forest leads to a distinction between two broad types of forest, primary or old-growth forest and secondary forest. There are also many natural factors that can cause changes in forests over time including forest fires, insects, diseases, weather, competition between species, etc. In 1997, the World Resources Institute recorded that only 20% of the world's original forests remained in large intact tracts of undisturbed forest. More than 75% of these intact forests lie in three countries—the boreal forests of Russia and Canada and the rainforest of Brazil.

In 2010, the United Nations Food and Agriculture Organization reported that world deforestation, mainly the conversion of tropical forests to agricultural land, had decreased over the past ten years but still continues at a high rate in many countries. Globally, around 13 million hectares of forests were converted to other uses or lost through natural causes each year between 2000 and 2010 as compared to around 16 million hectares per year during the 1990s. The study covered 233 countries and areas. Brazil and Indonesia, which had the highest loss of forests in the 1990s, have significantly reduced their deforestation rates. China instituted a ban on logging, beginning in 1998, due to the erosion and flooding that it caused. In addition, ambitious tree planting programmes in countries such as China, India, the United States and Vietnam – combined with natural expansion of forests in some regions – have added more than seven million hectares of new forests annually. As a result, the net loss of forest area was reduced to 5.2 million hectares per year between 2000 and 2010, down from 8.3 million hectares annually in the 1990s. In 2015, a study for Nature Climate Change showed that the trend has recently been reversed, leading to an "overall gain" in global biomass and forests. This gain is due especially to reforestation in China and Russia. However new forests are not completely equivalent to old growth forests in terms of species diversity, resilience and carbon capture. On September 7, 2015, the Food and Agriculture Organization of the United Nations released a new study stating that, over the last 25 years, the global deforestation rate has decreased by 50% due to improved management of forests and greater government protection.

Smaller areas of woodland in cities may be managed as urban forestry, sometimes within public parks. These are often created for human benefits; Attention Restoration Theory argues that spending time in nature reduces stress and improves health, while forest schools and kindergartens help young people to develop social as well as scientific skills in forests. These typically need to be close to where the children live, for practical logistics.

Canada


Canada has about 4,020,000 square kilometres (1,550,000 sq mi) of forest land. More than 90% of forest land is publicly owned and about 50% of the total forest area is allocated for harvesting. These allocated areas are managed using the principles of sustainable forest management, which includes extensive consultation with local stakeholders. About eight percent of Canada’s forest is legally protected from resource development. Much more forest land—about 40 percent of the total forest land base—is subject to varying degrees of protection through processes such as integrated land use planning or defined management areas such as certified forests.

By December 2006, over 1,237,000 square kilometres of forest land in Canada (about half the global total) had been certified as being sustainably managed. Clearcutting, first used in the latter half of the 20th century, is less expensive, but devastating to the environment, and companies are required by law to ensure that harvested areas are adequately regenerated. Most Canadian provinces have regulations limiting the size of clear-cuts, although some older clear-cuts can range upwards of 110 square kilometres (27,000 acres) in size which was cut over several years.

Latvia

Latvian Pine Forest in Ķegums Municipality
 
Latvia has about 3,270,000 hectares (12,626 sq mi) of forest land which equates to 50.6% of Latvia's total area (24,938 sq mi). 1,510,000 hectares of forest land (46.3% of total forest land) is publicly owned and 1,750,000 hectares of forest land (53.7% of total forest land) is in private hands. Latvia's forests have been steadily increasing over the years which is in contrast to many other nations, mostly due to the forestation of land not used for agriculture. In 1935 there was only 1,757,000 hectares of forest, today this has increased by more than 150%. Birch is the most common tree at 28.2% followed by pine (26,9%), spruce (18.3%), grey alder (9.7%), aspen (8,0%), black alder (5.7%), oak/ash (1.2%) and finally hardwood making up the rest (2.0%).

United States

In the United States, most forests have historically been affected by humans to some degree, though in recent years improved forestry practices have helped regulate or moderate large scale or severe impacts. However, the United States Forest Service estimates a net loss of about 2 million hectares (4,942,000 acres) between 1997 and 2020; this estimate includes conversion of forest land to other uses, including urban and suburban development, as well as afforestation and natural reversion of abandoned crop and pasture land to forest. However, in many areas of the United States, the area of forest is stable or increasing, particularly in many northern states. The opposite problem from flooding has plagued national forests, with loggers complaining that a lack of thinning and proper forest management has resulted in large forest fires.

Chestnut blight

From Wikipedia, the free encyclopedia

Chestnut blight fungus
Chestnut blight.jpg
Cankers caused by the fungal infection cause the bark to split.
Scientific classification
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C. parasitica
Binomial name
Cryphonectria parasitica

The pathogenic fungus Cryphonectria parasitica (formerly Endothia parasitica) is a member of the Ascomycota (sac fungi) taxon. It is native to South East Asia and was introduced into Europe and North America in the 1900s. The fungus spread rapidly and caused significant tree loss in both regions.

Overview

Cryphonectria parasitica is a parasitic fungus of chestnut trees. This disease came to be known as chestnut blight. Naturally found in South East Asia, accidental introductions led to invasive populations of C. parasitica in North America and Europe. The fungal disease has had a devastating economic and social impact on communities in the eastern United States. In the first half of the 20th century it killed an estimated four billion trees. Less severe impacts have occurred in Europe due to widespread CHV1 hypovirulence. CHV1 is one of at least two viral pathogens that weaken the fungus through hypovirulence and helps trees survive.

The American Chestnut and American chinquapin are highly susceptible to chestnut blight. The European chestnut is also susceptible but due to widespread CHV1 hypoviruluence, blight-induced tree death is less common. The fungus can infect other tree species such as oaks, red maples, staghorn sumacs, and shagbark hickories. Once infected, these trees will also exhibit orange bark with cankers. However, they will not exhibit shoot die back and death of the main tree. Instead the pathogen can persist in trees, but the fungus will spore and so may infect other trees. The fungus is spread by wind-borne ascospores and, over a shorter distance, conidia distributed by rain-splash action. Infection is local in range, so some isolated American chestnuts survive where there is no other tree within 10 km (6.2 mi). The root collar and root system of the chestnut tree have some resistance to blight infection due to soil organisms adversely reacting to the fungus; consequently, a large number of small American chestnut trees still exist as shoots growing from existing root bases. However, these regrown shoots seldom reach the sexually reproductive stage before being killed by the fungus.

History

North American infection

The chestnut blight was accidentally introduced to North America around 1904 when Chryphonectria parasitica was introduced into the United States from Japanese nursery stock. It was first found in the chestnut trees on the grounds of the New York Zoological Garden (the "Bronx Zoo") by Herman W. Merkel, a forester at the zoo. In 1905, American mycologist William Murrill isolated and described the fungus responsible (which he named Diaporthe parasitica), and demonstrated by inoculation into healthy plants that the fungus caused the disease. By 1940, most mature American chestnut trees had been wiped out by the disease.

Infection of American chestnut trees with C. parasitica simultaneously appeared in numerous places on the East Coast, most likely from Castanea crenata, or Japanese chestnut, which had become popular imports. Japanese and some Chinese chestnut trees have some resistance to infection by C. parasitica: the infection usually does not kill these Asian chestnut species. Within 40 years the nearly four-billion-strong American chestnut population in North America was devastated. — only a few clumps of trees remained in Michigan, Wisconsin and the Pacific Northwest. Because of the disease, American chestnut wood almost disappeared from the market for decades, although it can still be obtained as reclaimed lumber.

It is estimated that in some places, such as the Appalachian Mountains, one in every four hardwoods was an American chestnut. Mature trees often grew straight and branch-free for 50 feet and could grow up to 100 feet tall with a trunk diameter of 14 feet at a few feet above ground level. The reddish-brown wood was lightweight, soft, easy to split, very resistant to decay; and it did not warp or shrink. For three centuries many barns and homes near the Appalachian Mountains were made from American chestnut. Because of its resistance to decay, industries sprang up throughout the region to use wood from the American chestnut for posts, poles, piling, railroad ties, and split-rail fences. Its straight-grained wood was ideal for building l furniture, and caskets as well. The fruit that fell to the ground was an important cash crop and food source. The bark and wood were rich in tannic acid, which also provided tannins for use in the tanning of leather. Many native animals fed on chestnuts, and chestnuts were used for livestock feed, which kept the cost of raising livestock from being prohibitive.

Efforts started in the 1930s and are still ongoing, in Massachusetts and many other places in the United States, to repopulate the country with chestnut trees. Surviving American chestnut trees are being bred for resistance to the blight, notably by The American Chestnut Foundation, which aims to reintroduce a blight-resistant American chestnut to its original forest range within the early decades of the 21st century. Japanese chestnut and Chinese chestnut, as well as Seguin's chestnut and Henry's chestnut—have been used in these breeding programs in the US to create disease-resistant hybrids with the American chestnut. It's important to realize, though, that even Chinese chestnut trees vary considerably in blight resistance. Some individuals are quite susceptible while others are essentially immune to the disease.

Hypovirulence is not widespread in the US and attempts to commercially introduce CHV1 have not been widely successful. Though CHV1 persists in the applied tree, it does not spread naturally as it does in Europe, preventing it from being an effective form of biocontrol.

European infection

Chestnut blight was first identified around Genoa in the 1938. This quickly spread and was identified in France in 1946, Switzerland in 1951 and in Greece in 1963. It has most recently been found in the UK. Due to genetic differences between the fungal populations, it is likely that a second introduction of chestnut blight occurred in Georgia and Azerbaijan in 1938. The fungal infections initially caused wide-spread tree death in Europe. However, in the early 1950s trees were identified in Italy that survived fungal infection. On these trees the fungus caused more superficial cankers, that appeared to be healing. The reduced infection was due to the presence of CHV1, an RNA virus that infects C. parasitica. CHV1 spread naturally throughout Europe but is also spread artificially as a biocontrol measure (particularly in France). CHV1 is currently not present in the UK, Northern France or Eastern Georgia but introduction for biocontrol is being considered.

Symptoms

A chestnut tree that has been cut down, with blight on its inner bark and trunk
 
The fungus enters through wounds on susceptible trees and grows in and beneath the bark, eventually killing the cambium all the way round the twig, branch or trunk. The first symptom of C. parasitica infection is a small orange-brown area on the tree bark. A sunken canker then forms as the mycelial fan spreads under the bark. As the hyphae spread, they produce several toxic compounds, the most notable of which is oxalic acid. This acid lowers the pH of the infected tissue from around the normal 5.5 to approximately 2.8, which is toxic to plant cells. The canker eventually girdles the tree, killing everything above it. Distinctive yellow tendrils (cirrhi) of conidia can be seen extruding in wet weather.

Environment and disease cycle

The primary plant tissues targeted by C. parasitica are the inner bark, an area containing the conductive tissue, and the cambium, a layer of actively dividing cells that give rise to secondary vascular tissues. In these tissues, the pathogen forms diffuse cankers in which the mycelium overwinters. In the following spring, two types of fruiting bodies will form: pycnidia, usually first, and perithecia. Following rainfall, the pycnidia ooze orange tendrils of conidia, the asexual spores, while perithecia forcibly eject ascospores, the sexual spores. Upon becoming airborne, ascospores are carried by eddies of wind to new hosts or infect other parts of the same tree. When insects, birds, or other wild life come into contact with the cankers, they can mechanically disperse the conidia to a new host. Additionally, the asexual spores can be dispersed by rain splash. Once on the new host, or new area of the tree, the spores can germinate and infect the innerbark through insect wounds and fissures in the outer bark. 

If cankers continue to form and expand, the fungus can girdle the stem, severing the flow of nutrients and water to the vital vegetative tissues. The absence of nutrient dispersal will result in tree death, however, the root system will survive. As a result, American chestnuts exist mainly as shrubs sprouting from the old, surviving roots. However, these sprouts usually succumb to infection by C. parasitica before reaching sexual maturity.

Management: hypovirulence, sanitation, and chemical control

In Europe during the late 1960s, it was found that a strain of C. parasitica was less virulent, only able to produce shallow cankers that the tree could eventually form callus tissue over. The trait of hypovirulence could be transferred from an avirulent strain to a lethal strain through anastomosis, the fusion of hyphae. It was later discovered that this attenuated virulence was due to infection by a dsRNA mycovirus, Cryphonectria hypovirus 1 (CHV1).

Considering the nature of hypovirulent strains, there has been a strong interest to use them to manage lethal C. parasitica strains. In Europe, natural dissemination of hypovirulence in pathogen populations resulted in the restoration of economically valuable chestnuts. Unfortunately, this was not the case in the United States. Unlike Europe, the US has a greater diversity of C. parasitica strains. Thus, the spread of the mycovirus in American C. parasitica populations are inhibited by vegetative incompatibility, an allorecognition system that inhibits the fusion of hyphae between individuals that are genetically distinct at specific loci. Recently, however, “super mycovirus donor strains” of C. parasitica have been engineered to overcome this incompatibility system and could potentially be employed as a method of biological control.

In addition to biocontrol, chestnut blight can also be managed by sanitation practices and chemical control; however, such management strategies are only feasible on a small scale, such as in an orchard. Sanitation practices like the pruning of symptomatic limbs and removal of infected trees can serve to eliminate sources of inoculum and limit the spread of the pathogen. Additionally, some fungicides have been shown to be effective at controlling disease. In a study on the chemical control of chestnut blight in Castanea sativa, it was found that the external application of both copper oxychloride and carbendazim could reduce the rate of disease by almost 50%.

Conservation efforts in North America

There are approximately 2,500 chestnut trees growing on 60 acres near West Salem, Wisconsin, which is the world's largest remaining stand of American chestnut. These trees are the descendants of those planted by Martin Hicks, an early settler in the area. In the late 1800s, Hicks planted fewer than a dozen chestnuts. Planted outside the natural range of American chestnut, these trees escaped the initial wave of infection by chestnut blight, but in 1987, scientists found blight also in this stand. Scientists are working to try to save the trees. There is a program to bring American chestnut back to the Eastern forest and funded by the American Chestnut Foundation, Wisconsin Department of Natural Resources, USDA Forest Service, West Virginia University, Michigan State University, and Cornell University.

Removing blighted trees to control the disease was first attempted when the blight was discovered, but this proved to be an ineffective solution. Scientists then set out to introduce a hyperparasitic hypovirus into the chestnut blight fungus. The trees infected with virus-treated fungus responded immediately and began to heal over their cankers. However, the virus was so efficient at attenuating fungal growth that it prevented spreading of the virus from an infected fungus growing on one tree to that growing on another tree. Only the virus-treated trees recovered. Scientific opinion regarding the future of the stand varies.

Hybrid chestnut trees

Current efforts are under way by the Forest Health Initiative to use modern breeding techniques and genetic engineering to create resistant tree strains, with contributions from SUNY College of Environmental Science and Forestry, Penn State, the University of Georgia, and the US Forest Service. One of the most successful methods of breeding is to create a back cross of a resistant species (such as one from China or Japan) and American chestnut. Researchers identified two or three genes that allow for blight resistance, and are focusing on giving the American chestnut hybrids only those genes from the Chinese or Japanese chestnut.

The two species are first bred to create a 50/50 hybrid. After three back crosses with American chestnut, the remaining genome is approximate 1/16 that of the resistant tree, and 15/16 American. The strategy is to select blight-resistance genes during the back crossing, while preserving the more wild-type traits of American chestnut as the dominant phenotype. Thus, the newly bred hybrid chestnut trees should reach the same heights as the original American chestnut. Many of these 15/16 American chestnut hybrids have been planted along the East Coast, including in the Jefferson National Forest and on the Flight 93 National Memorial. Some of these sites have had researchers check on the saplings that have been planted to see their survival rate. For the hybrids to do well, they need areas with decent drainage and abundant sunlight. Meeting these needs can be hard to do, so not all restoration areas have been successful with hybrid survival.

Transgenic blight-resistant chestnut trees

Plant pathologists, Drs. William Powell and Charles Maynard, working at the State University of New York College of Environmental Science and Forestry, have developed American chestnuts which have full blight resistance. Full resistance was attained by introducing a wheat gene coding for the enzyme oxalate oxidase into the American chestnut genome. This enzyme breaks down the oxalic acid secreted by the fungus into carbon dioxide and hydrogen peroxide. Early studies on hypovirulence showed that less virulent strains of the chestnut blight produced less oxalic acid when attacking the cambium. The transgenic trees have blight resistance either equal to or surpassing that of Chinese chestnuts. In 2013, SUNY ESF had over 100 individual events being tested, with more than 400 slated to be in the field or in the lab for various assay tests in the next several years and more than 1,000 trees growing in several field sites in 2014. Government approval will be required before returning any of these blight resistant trees to the wild. The New York Botanical Garden has planted several of the transgenic trees for public display.

Economic and ecological impact of disease

In less than fifty years after its emergence, C. parastica virtually eliminated American chestnut as a canopy species in 8.8 million acres (3.6×106 ha) acres of forest. The chestnut fruit was a major food source for animals in the low elevation Appalachian forests. This loss resulted in a drastic decrease of the squirrel population, the extinction of seven native moth species, and the slowed recovery of deer, Cooper’s hawk, cougar, and bobcat populations. The effects of this disease also rippled further through the ecosystem, being linked to a decrease in the abundance of cavity-nesting birds and to a decrease in river water quality which negatively affected aquatic invertebrate populations.

In 1912, standing chestnut timber in just three states was estimated to be $82.5 million ($1.9 billion in current dollars) in value. Therefore, in addition to ecological impacts, C. parasitica potentially caused a devastating loss in economic welfare for communities dependent on the chestnut tree. Mountaineers, residents of Appalachian Mountain communities, had to drastically alter their life styles to cope with the effects of this disease.

Economic effects have also been considerable in Europe, particularly before CHV1 spreads naturally to a region. In Greece for example, the disease forced the migration of people who could not longer afford to live off chestnut trees. It has also lead to a 40% decline in Greek chestnut production.

Representation of a Lie group

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