Glomalin and Conservation in Humboldt County The 1996 discovery of the soil glue glomalin is changing our understanding of the impact of elevated carbon dioxide, while giving important clues to forest health, watersheds, revegetation, wildfire and carbon sequestration. Here I share what I have found so others may read and draw their own conclusions, and relate it to my own experience, Humboldt County issues and stories from the news.

Wednesday, July 14, 2004

59. Trinity River Flows Restored 

59. Trinity River Flows Restored
The San Francisco-based U.S. 9th Circuit Court of Appeals ruled that Trinity River must be allocated for fish twenty years after Congress mandated restoration of the river and its salmon. Agricultural, power and urban users were upset as much as 9% of currently appropriated water would be sent down the Trinity system to the Klamath for fisheries habitat restoration. Judges sounded angry it had taken twenty years for the water to start flowing in sufficient quantities as ordered by Congress and another ten since Congress ordered larger fish producing flows. Major fish kills have occurred on the Klamath downstream from the Trinity in the last few years, water conditions have been shown to be the cause.
This is clearly a case of those who grabbed something being unwilling to part with enough of it to allow natural processes to occur. The Yuroks are not about to be compensated for their losses over a century. No account is made of loss of commercial or sports fishing revenue. The claim of vital energy production falls flat, generating 1% of the State’s need. Agriculture was not considered by federal law in the first place and so had no say in the case.
In general, it is time to wake up and smell the coffee for those who are comfortable with the way things are despite insults to our natural systems. Agriculture can greatly reduce its water needs by using drip irrigation, glomalin accruing no-till farming, and better on site retention. A lot more water is lost to evaporation because large bodies of still water in high summer have large evaporation rates. Underground storage and shaded rivers reduce this loss.
Dams have been shown to interfere with fish passage, change water temperatures and eventually fill in with sediment, creating more problems. Each dam seems to have its own special interest group afraid of its losses , even where the dam is illegal or the current use is not what the dam was chartered for.
Groups opposed to dams have been able to accumulate large amounts of data showing the problems with dams on rivers throughout the country and the world. It is time for new thinking on every aspect of precipitation capture and storage, particularly in built up areas with large amounts of water absorbing land roofed, paved and landscaped into net water users rather than producers.
http://www.latimes.com/news/science/environment/la-me-trinity14jul14,1,1421464.story?coll=la-news-environment
Court OKs Trinity River Plan
The ruling upholds a federal order to increase flows to restore salmon habitat. It reduces supplies to farmers.
From Associated Press, July 14, 2004
SAN FRANCISCO — A federal appellate court approved a congressional plan Tuesday to increase flows into the Trinity River to restore fish habitat, reducing water to California farmers and hydroelectric plants. Most of the water in the Trinity, which originates in Northern California's Trinity Alps and flows west into the Klamath River, has been diverted for decades to service a fast-growing state where much of the water is located far from where people live and farm.
In 1984, Congress mandated restoration of the 112-mile-long Trinity River to combat dwindling supplies of salmon, steelhead and other aquatic life. In 2000, after years of studies, the Department of the Interior approved a plan to increase Trinity water. The plan was backed by Indian tribes who use the waters for sustenance fishing, while farming and hydroelectric power interests opposed it. The Trinity is a major artery in the Central Valley Project's system of dams, tunnels, canals and reservoirs that supply 200 water districts serving 30 million people in the agricultural rich Central Valley. It churns turbines for nine power generating stations.
The plan approved Tuesday diverts as much as 9% of the water project's capacity, depending on rain and snow amounts. The utilities argued the Interior Department's plan would decrease water flows that eventually reach the Central Valley, and that the government did not study the impact it would have on the millions of water users downstream. A spokesman for 600 California agricultural customers said farmers would probably get less water under the plan.
"That's water that is all part of a flow regime that is an important part of this large, complex interconnected water system," said Tupper Hull of the Westlands Water District, a Fresno-based agricultural water supplier that challenged the plan. A spokesman for the U.S. Bureau of Reclamation, Jeff McCracken, said the government did not study what, if any, impact the plan would have on farming because the law did not require it. "It's a fairly significant yield of water out of the system," McCracken said. "If there were an endless supply, this wouldn't have gone to court."
Westlands is considering asking the San Francisco-based U.S. 9th Circuit Court of Appeals to reconsider its ruling, Hull said. Hydroelectric utilities contended the government should further study the effect on energy production in light of California's energy crisis.
A three-judge 9th Circuit panel, however, was not persuaded, and reversed a lower court ruling that halted portions of the plan. The unanimous court said it was time to complete the "flow plan for the Trinity River." "Twenty years have passed since Congress passed the first major act calling for restoration of the Trinity River and rehabilitation of its fish populations, and almost another decade has elapsed since Congress set a minimum flow level for the river to force rehabilitative action," Judge Alfred Goodwin wrote.
Goodwin said less than 1% of California's energy production could be undermined. The Yurok Tribe, which straddles the Klamath in Humboldt and Del Norte counties downstream of the Trinity before it drains into the Pacific Ocean, celebrated the decision. The state's poorest tribe, which fishes the river for a subsistence living, was hit hard in 2002 when thousands of salmon died because of low flows. "The fish that use the Klamath also spawn in the Trinity. So a healthy Trinity River is important to a healthy Klamath River," the tribe's attorney, Scott Williams, said.
In the 1800s, Williams said, the 5,000-member tribe gave up thousands of acres in exchange for a promise its fishing would be protected. "It's been decimated by decades of dams, logging and diversions," said Williams, adding that the decision is a move "toward repairing that broken promise."
The plan calls for diverting 368,900 acre-feet of water to 815,200 acre-feet a year, depending on precipitation. Flows would be released from the Trinity Dam at different rates throughout the year to mimic natural flows. An acre-foot of water is enough to cover an acre of land to a depth of one foot, and contains 325,821 gallons, enough to supply one or two families for a year. The California Farm Bureau did not immediately comment on the decision.

Tuesday, July 13, 2004

58 Creating Wilderness 

Wilderness areas in Northern California may be increased by as much as 300,000 acres as a scheduled committee hearing on the proposal gets under way July 21 before the Senate Energy and Natural Resources Committee. Most of the land would add to existing wilderness areas. Our old foes, off-roaders and mountain bikers, are upset as usual at exclusion. They constantly ignore their impact on forest floors and drainages. There is some money for inholder acquisition on Federal lands. We can see line by l9ine discussions occurring on specific roads. We cannot see the reasoning behind wilderness areas needing to be “untrammeled by man”. Much existing wilderness is recovering lands left alone after overexploitation. People are everywhere. There is no vegetated place left without human impact. The twenty-first century definition of wilderness will need to add recovered lands as wilderness, while recreational users and commercial users will have to partner up on land use issues that negatively impact local environments, using production areas for multiple human uses like in Maine’s back country, where operating timberlands are closed but reopened for public recreation when a logging cycle is finished in that area.
Mountain bikers need to know bikes cause serious disruption to the mechanisms that prevent erosion, fungal networks of glomalin producing hyphae. They then can reasonably discuss options for the benefit of the ecosystem first and personal human pleasure in agreement with those basic principles of forest health.
Several years ago I asked BLM ntpo looking at condors being reintroduced to King Range. I thought there was pretty much land available for them. I learned condors from Big Sur could fly here as part of an average day. More and more problems were pointed out between people and condors. This area, in fact, is not that large to a condor, and the people have many problems with condors. Several were shot just last year.
Humans and humans reacting to threats to their economies are the major cause of conflict. Unlimited human growth will continue to press species and landscapes across the globe. Creating wilderness hedges the bet for future generations.

http://www.times-standard.com/cda/article/print/0,1674,127%257E2896%257E2269473,00.html
Wilderness bill will go to key U.S. Senate committee Tuesday, July 13, 2004 -
John Driscoll
The Times-Standard
A bill that would mark 300,000 acres of public land in Northern California as wilderness will get a hearing before a Senate committee this month.
Wilderness advocates say they're enthusiastic about the Senate Energy and Natural Resources Committee hearing July 21. They say the bill enjoys broad support, and that the legislation will protect vital and beautiful areas from logging, mining and off-road vehicle use.
But there is dissent, not surprisingly, from four wheelers and some mountain bikers. They claim the bill would close some key recreational roads, despite staunch insistence otherwise from proponents.
The Northern California Coastal Wild Heritage Wilderness Act in most case adds acreage to existing wilderness areas. There is some language to provide money to acquire inholdings on federal land.
One of several wilderness bills slogging through the Legislature, this one deals with areas in Rep. Mike Thompson's 1st Congressional District.
They include additions to the Siskiyou, Trinity and Yolla Bolly wildernesses, an area called Mad River Buttes south of Titlow Hill, and a 42,000-acre area in the King Range National Conservation Area.
Josh Buswell-Charkow of the California Wilderness Coalition said some of the areas face threats from illegal off-road vehicle use and logging.
Buswell-Charkow said his group went to great lengths to make sure no legal roads are closed through a wilderness designation. Motorized vehicles and mountain bikes are not allowed in wilderness areas. Hikers and horseback riders are.
Don Amador of the Blue Ribbon Coalition in Idaho hotly disputes that. He said that part of the Smith-Etter Road in the King Range would be closed, and that some spur roads used by hunters in Six Rivers National Forest would also be shut off.
He also said some of the areas don't fit the "untrammeled by man" description lined out in the 1964 Wilderness Act. Amador showed photos of radio towers, old asphalt roads and logging in the proposed wildernesses.
"They're trying to create a wilderness for the 20th century that in no way resembles the provision of the 1964 act," Amador said.
Thompson emphatically said that no legal roads will be closed in his bill.
"These are exceptional properties and they deserve the enhanced status of wilderness," the St. Helena Democrat said.
The bill also allows the U.S. Bureau of Land Management and the U.S. Forest Service to use any means necessary to fight fires in the areas.
The bill contains appropriations of $1.25 million for three years for restoration and $23 million a year for law enforcement, acquiring inholdings, fire fighting and tourism development.
For some, there is inner conflict over the bill. Justin Brown, co-owner of Revolution Bicycle Repair in Arcata said he is generally concerned about losing areas to ride. He said that the closer one gets to the San Francisco Bay area, the fewer options exist for backcountry riding.
But, Brown said, he's not opposed to wilderness designation if it stops commercial uses of the areas.
"I love to mountain bike and I would love access to that land to ride on," Brown said. "But I don't want to see that area logged over."

57. Rolling Back the Rules on Roads and Condors 

57. Rolling Back the Rules on Roads and Condors Two important articles today demonstrate the problems conservationists deal with on a regular basis. These problems go beyond individual sites or rules and are much more related to the old development versus nature debates. Unfortunately, the developers have won so many times conservationists are often content with whatever mitigation operators are willing to throw them. When the mitigations are challenged by development or industry, the original zeal is often gone and huge sums of taxpayer money have gone to subvert those protections. In the first article, no mention is made of the now well documented destruction of forests and streams as a result of road building, which inspired the rule in the first place. Forest road building the traditional way has had major negative impacts on fisheries.
Nowhere in the article or the many road workshops I have attended is there anything like the erosion control measures on building sites back East demonstrated in Erosion Control magazine. Storm water is filtered, silt fencing surrounds all impact sites, landscape material and coir rolls protect stream banks from muddy runoff, products that bind soil particles, allow infiltration of road surfaces are coming online. These practices may be too expensive for profitable operation but they surely will evolve into more environmentally and developer friendly methods. But at this point, they want to be able to work as cheaply and quickly as possible. Because whatever rules have been put n place are inadequate or stop development entirely, results are uneven and no agreement has been reached.
In the second article we revisit old friends HCP and “careful what you wish for” on one of the largest subdivisions in California in an area where condors have been released. Habitat Conservation Plans supercede state regulations allowing some impacts to ESA listed species if the work improves or expands critical habitat, often improved as mitigation for development. Usually you get a protected percentage of a locale with some modification related to the local issue for the species benefit.
It is amazing how words can change meaning in their use. Condors are describes in a negative way as large, far-ranging, curious, needing large areas of undeveloped country and sometimes attracted to housing areas. All the effort to find habitat, capture the birds, breed them, find suitable habitat for reintroduction at thirty million bucks seems like a bargain for seventeen years of effort, but developers will claim they have lost more than that in mitigations and restrictions. They have been planning for ten years too.
As always in development cases, the money and lasting power of corporations is used to continually revisit a subject until the opposition fails. This is story of the ban on off-shore oil drilling in California., port development, water sales out of the area and a host of other issues. As the level of awareness rises, we can expect more battles to take place. While the exact issues and outcomes will vary, the results will be more or less impaired natural systems slowly recovering from massive damages in the name of human progress.
Witness the problems generated by damming rivers as one example we are all aware of. Does this mean we can remove old and less than useful dams? No. We can’t even prevent them from getting renewal permits on a fifty year basis. Are they impacting ESA listed species? Absolutely. When the chartered reason for their existence is questioned the claim of public need and grand fathered uses overwhelms obvious common sense. What weighs the difference? Perceived loss of doing things the way folks are used to with the cash flowing in the usual swales.
Cash is like rainfall in an unstable but recently roaded area. When it pours, all the old drainages are overwhelmed. Old lines of usual activity are blurred and new wealth generated, adding voices to the development side and further drowning out criticism. The new voices attack defenders of natural systems as costing them money and opportunity. The defenders say the natural world will eventually prevail- it always does. It is up to us to learn to live inside these sustainable natural systems, or nature will replace us with species that can and will do just that.
Humans are not the only species degrading pre-existing communities. Over and over, alien species are setting up shop in natural areas threatening to forever change species makeup and populations of large numbers of plants and animals. IN nearly every case, we have a less valuable invader without local enemies grabbing up habitat at amazing rates and impacting other ecosystem residents. There may be some convergence of biological operation at work. For example, shellfish that filter feed are a necessity for cleaning waters in bays and harbors. Native species, small and uneconomical, were the natural species. In some areas, East Coast oysters, which grow larger and have better market appeal, replaced them. The natives were over fished initially and then not favored, thus declining in importance to the health of the ecosystem. Now the exotic zebra mussel has arrived and has had a tremendous impact. They are crowding out regular inhabitants, grow on anything in the water, damaging boats, sluice gates, filter screens at power plants and breeding grounds of other species.
Nature will always fill a vacuum. The one constant in the world is change. Human activity always degrades natural systems. Humans will eventually go back to living in the landscape and its natural laws. In the meantime, we have become the third largest biomass of any individual species, impacting every natural system on earth.
http://www.latimes.com/news/science/environment/la-na-roadless13jul13,0,6775857.story?coll=la-news-environment New Forest Rules May Pave Way for Roads
Bush plan would sweep aside Clinton policy that protected 58.5 million acres of federal land.
By Bettina Boxall, LA Times Staff Writer, July 13, 2004
The Bush administration proposed new forest rules Monday that could lead to logging, mining and oil and gas development in remote country that had been protected under a policy issued in the waning days of the Clinton presidency.
The regulations would replace a January 2001 rule that banned road building and timber cutting on 58.5 million acres of roadless terrain in national forests with a policy giving state governors a say in the backcountry's management. Most of the land is in 12 Western states, including 4 million acres in California.
Hailed by conservationists, the road prohibition was quickly challenged in a series of lawsuits filed by states and various interest groups that complained it was creating de facto wilderness areas, usurping congressional authority. Early court decisions were conflicting, with two federal district judges ruling against the Clinton road ban and a federal appeals court upholding it.
The Bush administration proposal, announced in Boise, Idaho, by Agriculture Secretary Ann M. Veneman, would give governors considerable input on the future of roadless areas. States could petition the federal government if they wanted to maintain road-building bans on all or part of the affected forestland. They also could ask federal officials to open the land to road construction, whether for logging, gas or oil development or off-road vehicle use. The decision on any petition would be made by the Agriculture secretary.
Mark E. Rey, the Agriculture undersecretary who oversees the U.S. Forest Service, said the proposed regulations were an attempt to resolve a 40-year fight over roadless areas, which make up about 30% of the country's national forests. Broad, sweeping policies, such as that issued by Clinton, haven't worked, nor have attempts to settle the issue on a forest-by-forest basis, Rey said.
"We hope that this is a middle way or a third way involving the governors to do two things," Rey said. "One is to bring good site-specific information and scientific data, and to bring to bear enough political closure to get people to agree."
He predicted the new rule's result would not be that different from the Clinton policy. "Are we going to be seeing a significant modification of the Clinton roadless rules? I don't think so," Rey said.
Among those applauding the new policy were the chairman of the House Resources Committee, Republican U.S. Rep. Richard W. Pombo of Tracy, who sees it as a welcome departure from the Clinton rule.
"This proposal embraces the fact that local people are the best stewards of our forests," Pombo said. "It injects common sense and local control into Clinton's 11th-hour, mindless edict. Forest management decisions should be made at the state level by people who know individual forest conditions best, not by bureaucrats surrounded by concrete in Washington. Today's decision will be praised by Americans throughout the West, where 90% of these roadless areas occur."
Michael Mortimer, chairman of the Society of American Foresters policy committee, said his organization was generally pleased with the new approach. "We think it takes into account the regional difference in states and gives land managers more flexibility," he said.
The Bush proposal was criticized by groups as diverse as Taxpayers for Common Sense and the Outdoor Industry Assn., a trade group for manufacturers and retailers of outdoors equipment and clothing.
"This 'opt in' approach to roadless management provides no guarantees of real, long-term protections for roadless areas," said Frank Hugelmeyer, president of the outdoor association. "The future of recreation destinations essential to the health of the outdoor recreation industry is at stake."
Without networks of roads, or the logging or mining that can go with it, the roadless forests are prized by conservationists as places of quiet and natural beauty where hikers or hunters can escape in solitude.
Doug Honnold, an attorney for the environmental group Earthjustice, who has defended the roadless rule in some of the nine lawsuits filed against it, said the proposed regulations opened the door to industrial development in the backcountry.
"The state governors can request more logging and more road building and more oil and gas development and more hard-rock mines than we've ever had before in these areas," Honnold said. "If your goal is to maximize the amount of corporate development of our national forests, this is a great plan. If your goal is to protect clean drinking water and wildlife, it's an awful plan."
Although about a fifth of California's national forestlands have no roads, the proposed regulations will probably have a greater effect in other Western states. That is because of the slightly more than 4 million roadless acres in the state, only about 400,000 are considered suitable for timber production, according to Matt Mathes, a Forest Service spokesman in California.
"We have been staying out of our roadless areas for quite some time in California, and we have no plans to build roads in roadless areas in California," Mathes said. "There is nothing on the books for the foreseeable future."
A spokeswoman for Gov. Arnold Schwarzenegger said he welcomed the chance to be involved but had to review the proposal. "The governor agrees it's important for states to be a part of this process," said spokeswoman Ashley Snee.
Pointing out that most of the potential oil and gas fields in the Los Padres National Forest in Central and Southern California lie within roadless areas, Sara Barth, California director of the Wilderness Society, predicted the new policy would open up remote areas in the state.
"Roads are the access point for all kinds of development that winds up harming the forests," she said.
Giving state governors so much potential sway in the management of federal forests represents a dramatic departure from past practice, said Sean Hecht, executive director of the UCLA Environmental Law Center.
The government will take comments on the proposal for the next 60 days before issuing a final
rule.

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http://www.latimes.com/news/science/environment/la-me-condor13jul13,1,2363064.story?coll=la-news-environment
Developer Wants to Ease Condor Rules
Tejon Ranch asks to be shielded from penalties if birds are hurt during building of homes.
By Daryl Kelley, LA Times Staff Writer, July 13, 2004
The Tejon Ranch Co. has applied for a federal permit to protect developers if they accidentally harm or kill the endangered California condor by building three projects on the vast ranch north of Los Angeles.
Tejon Ranch is seeking an "incidental take" permit, which would allow the developer to "harm, harass, trap, shoot or kill" North America's largest bird during or after planned construction of a 23,000-home residential project, a 1,450-acre warehouse park and a mountain resort community along Interstate 5 in the Tehachapi Mountains.
Ranch officials and the U.S. Fish and Wildlife Service are set to jointly explain the proposal at a public meeting in Frazier Park today. The permit process could be complete by late this year, officials said.
"The misconception is that this permit would allow for the killing of the bird," said biologist Rick Farris, a division chief of the U.S. Fish and Wildlife Service in Ventura. "What this permit does is allow them to conduct their lawful activities and assure that what they're doing is not going to cause the extinction of the species."
If the notoriously curious and sometimes destructive condors are drawn to the new developments near their Tejon habitat, ranch officials and the wildlife service have crafted plans to deal with them, Farris said. They include having a biologist on the ranch to frighten away the birds if they land on houses or swoop into backyards to swallow debris.
But environmental groups and local residents — notified of the proposal in a June 25 posting in the Federal Register — are rallying to oppose the permit. "Giving a permit to harass, harm or kill California condors really goes against all the work being done to turn around the fate of that species," said Kerri Camalo of Defenders of Wildlife, a national group. "It seems an affront to the people who have spent so much time and taxpayers' money to keep this species from becoming extinct," she said. "Now we'd allow the few condors alive in the wild to be taken in the name of development."
Ninety-nine of the rare condors live in the wild, including 49 in California, after a 25-year, $35-million effort to save the bird from extinction. An additional 149 are in captivity.
"Actually, I'm a little concerned about this," said Jesse Grantham, interim coordinator of the government's condor recovery program. "This is sort of a collision between wildlife and humans," he said. "How do we deal with an animal that can cover great distances and needs significant space to survive? The condor is a flagship species, and the flags are flying everywhere."
The 270,000-acre Tejon Ranch — a prized wildlife habitat that stretches 40 miles from north to south and 21 miles across — is a favorite feeding and resting spot for California condors. Indeed, the last condor taken into captivity, in 1987, in an effort to save the species was captured at Tejon Ranch. The birds now migrate from as far away as Big Sur and from nests in the mountains above Fillmore to hang out along wind-swept ridgelines at the southern end of the San Joaquin Valley. They love to soar with the afternoon zephyrs, experts say.
But just a few miles from those ridgelines, the Tejon Ranch resort community is planned. Authorities fear it would attract the big birds because of its high elevation in the mountains that make up the ranch's rugged backbone. Tejon Ranch general counsel Dennis Mullins said Monday that the company had worked with federal officials for more than 10 years to come up with ways to assure that condors and the development can safely coexist.
"We're going to do good things for the condor," Mullins said. "We asked in 1992, when the condors were being reintroduced to the wild, how Tejon Ranch could help the condor recovery program without impairing its property rights."
Federal officials and the development firm came up with a set of guidelines for where and how dwellings can be built, and a series of options on how to deal with condor problems if they develop. Tejon Ranch has agreed with recommendations to ban buildings along ridgelines, because they would attract the birds. Strict limits would also be placed on height for the same reason. And the design of backyards would be altered so they won't lure the big birds.
"But there is the potential that condors will start hanging around the houses; they've done it in the past," said Farris of the Fish and Wildlife Service. "When they do that we have to change their behavior. That may even involve removing them from the wild and putting them back in captivity. Although it wouldn't necessarily kill them, we're saying they would be ecologically dead in terms of their ability to contribute to the population."
This afternoon's meetings — at 3 p.m. and 7 p.m. at Cuddy Hall in Frazier Park — are the first public parts of an environmental review process that Farris said he began in 1998, and will lead to the release this fall of two related analyses on the effect of development on Tejon Ranch.
The "incidental take" permit is a required part of a so-called habitat conservation plan being prepared jointly by Tejon Ranch and the Fish and Wildlife Service. The federal Endangered Species Act requires an evaluation of a project if the development site is important as a habitat of an endangered species. Because of its size, location and variety of wildlife, many experts consider Tejon Ranch as perhaps the most environmentally important undeveloped stretch of land in California. Ranch owners offered last year to sell 100,000 acres to an environmental trust, a proposal that is still pending. "We have a responsibility to process this [application] to the best of our ability," Farris said. "At a later time, we'll make a decision on whether this is something we should be doing, or not."

56. Smoke Chemical Shown to Increase Germination 

56. Smoke Chemical Shown to Increase Germination
Science continues to improve our understanding of natural phenomena and give us tools for manipulating natural processes. This is particularly interesting in helping hard to regenerate native plants for restoration and natural landscaping. The article seems to imply this is an easily manufactured product we may see in the stores soon.
http://www.latimes.com/news/science/environment/la-sci-smoke10jul10,1,6814912.story?coll=la-news-environment
Smoke Chemical Shown to Increase Germination
By Eric D. Tytell
Los Angeles Times Staff Writer
July 10, 2004

Plant biologists, who have long known that a compound in smoke causes many seeds to sprout, have now isolated the specific chemical — a finding that could help boost crop yields and preserve rare plants.
The study, published Friday in the journal Science, describes the structure and synthesis of a compound called a butenolide, a component of smoke.
Scientists at the University of Western Australia tested the chemical on seeds from lettuce, tobacco and 14 wild plants. They showed that it dramatically increased the number of seeds that sprouted.
The chemical could help biologists cultivate endangered plants that require fire to sprout — without resorting to actual fires. It could also make farming more cost-effective because the compound increases the germination rate for many crops, even if they do not require fire.
"There's potentially a whole new way that everyone — from the botanist to the vegetable grower — could get a benefit," said Kingsley W. Dixon, science director at Kings Park and Botanic Garden in Perth, Australia, and a coauthor of the study.
Biologists think many seeds evolved to respond to fire, which might signal the availability of more space and light to grow. Some seeds, like those from sequoias, almost never sprout without this signal.
"The real power of this discovery is going to be in the management of wild lands," Dixon said.
Seeds from species like lettuce are also sensitive to fires and germinate about twice as often when watered with even low concentrations of the chemical, the researchers found.


Monday, July 12, 2004

55. Organic Farming Sequesters Atmospheric Carbon and Nutrients in Soils 

These are the findings of a twenty-three year study by Rodale Press (Organic Gardening)of carbon and nitrogen storage in soils. The article includes farming, forestry, numbers, glomalin, sequestration, water and is just chock full of good stuff.



Organic Farming Sequesters Atmospheric Carbon and Nutrients in Soils

Paul Hepperly, The New Farm® Research Manager
The Rodale Institute®

Executive Summary



Organic farming may be one of the most powerful tools in the fight against global warming. Findings from The Rodale Institute’s 23-year Farming Systems Trial® (FST) comparing organic and conventional cropping systems show organic/regenerative agriculture systems reduce carbon dioxide, a major greenhouse gases-positioning organic farming as a major player in efforts to slow climate change from runaway greenhouse gases increases.

Besides being a significant underutilized carbon sink, organic systems use about one third less fossil fuel energy than that used in the conventional corn/soybean cropping systems. According to studies of the FST in collaboration with Dr. David Pimentel of Cornell University, this translates to less greenhouse gases emissions as farmers shift to organic production. The ability of organic agriculture to be both a significant carbon sink and to be less dependent on fossil fuel inputs has long-term implications for global agriculture and its role in air quality policies and programs.

Since 1981, data from the Farming Systems Trial has revealed that soil under organic agriculture management can accumulate about 1,000 pounds of carbon per acre foot of soil each year. This accumulation is equal to about 3,500 pounds of carbon dioxide per acre taken from the air and sequestered into soil organic matter. When multiplied over the 160 million acres of corn and soybeans grown nationally, a potential for 580 billion pounds of excess carbon dioxide per year can be sequestered when farmers transition to organic grain systems.

It is believed that agricultural soil has a significant potential to capture and retain or sequester carbon dioxide. The 1995 Kyoto Protocol references this potential without emphasizing its capacity nor the importance of organic agriculture management for this purpose. Since then, researchers have moved forward strongly with investigations to support agriculture’s real potential to sequester carbon. The Rodale Institute® findings have taken this one step further by measuring carbon content and studying the positive impacts of carbon sequestration in organically-farmed soils.

The Rodale Institute’s 23-year findings show that organic grain production systems increase soil carbon 15 to 28%. Moreover, soil nitrogen in the organic systems increased 8 to 15%. The conventional system showed no significant increases in either soil carbon or nitrogen in the same time period. Soil carbon and nitrogen are major determinants of soil productivity.

Why does the soil carbon level increase in organic systems but not in conventional systems when crop biomass is so similar? We believe the answer lies in the different decay rates of soil organic matter under different management systems. In the conventional system the application of soluble nitrogen fertilizers stimulates more rapid and complete decay of organic matter sending carbon into the atmosphere instead of retaining it in the soil as the organic systems do.

Additionally, soil microbial activity, specifically the work of mychorrhiza fungi, plays an important role in helping conserve and slow down the decay of organic matter. Collaborative studies in our Farming Systems Trial® with the United States Department of Agriculture Research Service (ARS) researchers, led by Dr. David Douds, show that mychorriza fungi are more prevalent in the FST organic systems. These fungi work to conserve organic matter by aggregating organic matter with clay and minerals. In soil aggregates, carbon is more resistant to degradation than in free form and therefore more likely to be conserved. Support for this work comes from United States Department of Agriculture researchers at the Sustainable Agriculture Laboratory in Beltsville, Maryland. Their findings demonstrate that mychorrizal fungi produce a potent glue-like substance called glomalin that is crucial for maximizing soil aggregation. We believe that glomalin is an important component for carbon soil retention and encourage increased investigation of this mechanism in carbon sequestration.

Increasing soil organic matter for the soil’s carbon bank is a principle goal of organic agriculture. Organic agriculture relies on the carbon bank and stimulated soil microbial communities to increase soil fertility, improve plant health, and support competitive crop yields. This approach utilizes the natural carbon cycle to reduce the use of purchased synthetic inputs, increase energy resource efficiency, improve economic returns for farmers, and reduce toxic effects of fertilizers and pesticides on human health and the environment.

US Secretary of Agriculture, Ann Veneman, puts it this way, “The technologies and practices that reduce greenhouse gases emissions and increase carbon sequestration also address conservation objectives, such as improving water and air quality and enhancing wildlife habitat. This is good for the environment and good for agriculture.”




Organic Farming Sequesters Atmospheric Carbon and Nutrients in Soils


Background, Findings, and The Next Steps

An analysis of gases trapped within glacier ice shows that 18,000 years ago, during the last ice age, atmospheric concentrations of carbon dioxide were 60% lower than those found in the atmosphere today. This low concentration of carbon dioxide was associated with a 4° C (about 10° F) drop in average temperature. Presently, global atmospheric carbon dioxide levels are 25% higher than in the late 1800’s. If emissions continue at current levels, carbon dioxide in the atmosphere may double or even quadruple within the next 100- 300 years.

In 1938, G. Callendar published findings suggesting that the burning of fossil fuels, such as coal, oil and natural gases, would likely increase world temperatures. Since 1958, continuous carbon dioxide measurements on Mount Mauna Loa in Hawaii confirm that carbon dioxide is increasing in the atmosphere at a rate of about 1.3 parts per million (ppm) per year. Atmospheric scientists believe that although several other gases contribute to the greenhouse effect in the Earth’s atmosphere, carbon dioxide is responsible for over 80% of potential warming. NASA scientist James Hansen tracked temperature changes in relation to past carbon dioxide levels and he correlated the 25% increase in carbon dioxide over the last 100 years with a 0.7° C warming of the atmosphere. A number of models have predicted that at current rates of carbon dioxide emission, the Earth will warm 2.5° C in the next 100 years at current rates of carbon dioxide emission.

According to climatic change models, agriculture could be seriously affected by global warming. It is estimated that 20% of potential food crop production is lost each year due to unfavorable weather patterns (drought, flood, severe heat and cold, strong storms, etc.). The deterioration of weather patterns in North America could have devastating effects on world supplies of basic food grains such as wheat and corn. Climate change modelers predict that higher temperatures will generate more extreme weather events, such as severe droughts and torrential rains. A shift of 1 to 2° C in summer temperatures at pollination season can cause a loss of pollen viability, resulting in male sterility of many plant species such as oats and tomatoes.

As global temperatures rise, the glaciers and polar icecaps will melt, leading to major island- and coastal-flooding. About 50% of the United States population lives within 50 miles of a coastline. As coastlines move inland, uncontrolled carbon dioxide levels will directly affect coastal dwellers. If greenhouse gases continue to increase in the next several hundred years, the rise of global temperature is estimated at 7° C, or almost 15° F, and the sea level would rise over 2 meters, or in excess of 6 feet.


Soil Organic Matter-Key to Sequestration

Normal seasonal carbon dioxide fluctuations in the atmosphere demonstrate that plant growth governs major amounts of carbon dioxide, enough to change atmospheric concentration by up to 10 ppm. By increasing plant production, we can reduce carbon dioxide concentrations in the atmosphere. Carbon dioxide levels are minimized in summer when vegetation is lush, and maximized in winter when plants die or go dormant. The fluctuation of carbon dioxide from season to season (about 10 ppm) is about 7 times greater than the yearly average increase in atmospheric carbon from fossil fuel burning and deforestation (1.3 ppm). Plants serve as sinks for atmospheric carbon dioxide. Carbon stored in vegetation, soil, or the ocean, which is not readily released as carbon dioxide, is said to be sequestered. To balance the global carbon budget, we need to increase carbon sequestration and reduce carbon emissions. While carbon can cycle in and out of soil or biomass material, there are methods for building up what are called soil “humic” substances (also known as organic matter) that can remain as stable carbon compounds for thousands of years.

Before forests and grasslands were converted to field agriculture, soil organic matter generally composed 6 to 10% of the soil mass, well over the 1 to 3% levels typical of today’s agricultural field systems. The conversion of natural grasslands and forests around the globe works to elevate atmospheric carbon dioxide levels significantly. Building soil organic matter by better nurturing our forest and agricultural lands can capture this excess atmospheric carbon dioxide, and preserve more natural landscapes.

Agricultural and forest carbon sequestration will reduce the dangers that carbon dioxide currently presents to our atmosphere and world climatic patterns. These benefits will complement energy conservation and emission control efforts. Improved energy use is important because if all fossil fuel reserves were used in the next several hundred years, carbon dioxide in the atmosphere would increase 4 to 8 times present levels (currently the atmosphere holds 750 Gigatons of carbon, while known fossil fuel energy reserves hold 5,000 Gigatons of carbon.). Soil organic carbon, even at its present depleted level (1,580 Gigatons of carbon[C]), is still estimated to be almost double the quantity of all the carbon currently found in the atmosphere as carbon dioxide (800 Gigatons C), and about three times the amount found in all living organisms on the planet (500 Gigatons C).

Soil, agriculture, and forests are essential natural resources for sequestering runaway greenhouse gases helping to derail drastic climate changes. The amount of carbon in forests (610 Gigatons) is about 85% of the amount in the atmosphere. The 1998 Resources For the Future Climate Issue Brief #12 states, “Although it is well known that the world’s tropical forests are declining, it is less widely recognized that the world’s temperate and boreal forests have been expanding, albeit modestly…Nevertheless, overall, the size of the global forest carbon stock appears to be declining, thereby generating a net carbon source.”


The Rodale Institute Farming Systems Trial® Findings

Agriculture is, and always will be, a major tool in carbon sequestration. The Rodale Institute’s 23 year Farming Systems Trial® research provides real world experience and the starting point for understanding the potential for agriculture to reduce greenhouse gases. The FST® is the longest running agronomic experiment designed to compare organic and conventional farming methods and production systems.

Since 1981, The Rodale Institute® has continuously monitored soil carbon and nitrogen in its Farming Systems Trial® (FST). Carbon and nitrogen monitoring is just one component of a comprehensive battery of soil quality, economic and energy data that The Rodale Institute researchers gathered over the 23-year lifespan of the FST®. Researchers at The Rodale Institute believe that soil carbon and nitrogen findings are especially significant and dramatic. In the organic systems, soil carbon increased 15-28%, demonstrating the ability of the organic systems to sequester significant quantities of atmospheric carbon. Specifically, the FST organic manure system showed an average increase of soil carbon of about 1000 lbs per acre-foot of soil per year, or about 3,500 pounds of carbon dioxide per acre-ft per year sequestered. When multiplied over the 160 million acres of corn and soybeans that are produced nationally, a potential of an increase of 580 billion pounds of carbon dioxide per year would be sequestered by farmers switching from conventional chemically based farming systems to organic grain farming methods.

Additionally, in the organic systems, soil carbon has increased 15 to 28%. Over the 23 year lifespan of the FST, the conventional system showed no significant increases in either soil carbon or nitrogen. This demonstrates that organic farming methods increase stored carbon and retain other nutrients because organic soils hold these nutrients in place for uptake by plants. In the process, reduce nitrate and other nutrient runoff into streams and water aquifers. These findings can be beneficial to all farmers by helping them to increase crop yields while decreasing energy, fuel and irrigation costs.

We believe this is the longest scientifically replicated study that has been continuously monitored for soil quality including carbon and nitrogen levels. Certainly study is a first in terms of its duration and comparison of the carbon sink potential of organic and conventional agriculture soils. This study gives us a baseline for developing an ambitious scale of work to replicate and then accelerate the carbon sequestration potential of organic farming methodologies.

In addition to capturing more carbon as soil organic matter, organic agricultural production methods also emit less greenhouse gases through more efficient use of fuels. Energy analysis of The FST by Dr. David Pimentel from Cornell University show that organic systems use only 63% of the energy input used by the conventional corn and soybean production system. In all systems, yields of corn and soybean were not different, except in drought years, when organic systems yielded 25 to 75% more than the conventional system. The organic yield advantage in drought years is specifically related to the ability of higher-carbon organic soils to capture and deliver more water to crop plants. Dr. David Pimentel’s findings show that the biggest energetic input, by far, in the conventional corn and soybean system is nitrogen fertilizer for corn, followed by herbicides for both corn and soybean production.

Organic farming also makes economic sense. In addition to reducing input costs, economic analysis by Dr. James Hanson of the University of Maryland has shown that organic systems in the FST are competitive in returns with conventional corn and soybean farming, even without organic price premiums. Real world organic price premiums allow farmers to take advantage of certified organic production systems to achieve economically viable returns without massive governmental subsidies.

How can low input organic systems be competitive in productivity with a high input chemically based conventional system? USDA scientist, David Douds, in collaboration with scientists at The Rodale Institute®, has shown that in the organically managed systems, the biological support system of mycorrhiza fungi is much more robust and the fungi are more prevalent, active, and diverse. Synthetic chemical fertilizers and pesticides inhibit mycorrhizae. In organic production systems, increased mycorrhiza fungal activity allows plants to increase their access to soil resources, thereby stimulating plants to increase their nutrient uptake, water absorption, and their ability to suppress certain plant pathogens.

The process and ability of mycorrhiza to sequester carbon has perhaps an even greater significance. Mycorrhiza fungi produce a novel glue-like substance called glomalin. Glomalin stimulates increased aggregation of soil particles. Soil particle aggregation results in an increased ability for soil to retain carbon. The role of mycorrhiza and glomalin in soil carbon retention requires further investigation. Other biological mechanisms that will result in a greater ability of soil to sequester carbon naturally and to improve soil properties require further investigation as well.


Benefits Beyond Carbon Sequestration

The presence of sequestered carbon in The Rodale Institute’s FST® organic field trials is an indicator of healthy soil because healthy soil is abundant in carbonaceous matter, in particular the organic material humus. It is humus that enables healthy soils to retain water during periods of drought; as well as retaining mobile nutrients found in soils such as phosphates and nitrates, that would otherwise be lost as runoff to streams and aquifers.

These trials are illustrative of both economic benefit as well as environmental protection working hand in hand. The economic benefits are realized by farmers and landowners who see reduced costs for fertilizer, energy fuels and irrigation, and increased crop yields at the same time. It is also economically beneficial to the agricultural business economy, and an environmental benefit to all of us, that specific soil management and tillage practices can help to sequester or retain carbon in the soil--carbon that would otherwise be lost to the atmosphere as a component of greenhouse gases.

In summary, organic farming can reduce the output of carbon dioxide by 37-50%, reduce costs for the farmer, and increase our planet’s ability to positively absorb and utilize greenhouse gases. These methods maximize benefits for the individual farmer as well as for society as a whole. It is a winning strategy with multiple benefits and virtually no risk. These proven approaches mitigate current environmental damages and promote a cleaner and safer world for future generations.


The Next Steps

In recent months, staff from The Rodale Institute® met with officials of the Pennsylvania Departments of Agriculture and Environmental Protection. Together, we are working on a Statement of Cooperation that will provide a platform for future research and education on how organic farming can provide significant economic and environmental benefits. With 22 years of data from the FST® field trials in place, we will explore ways to promulgate and systematize the knowledge that has been gathered. In recent years, other researchers around the world have also begun to investigate and document the potential for soil carbon and nutrient sequestration. It is important to move forward quickly to lead the research in this field.

First, we propose to review the current body of scientific literature to determine if there are ways to accelerate the formation of organic material in soil, and to determine if it is possible to predict the rate of carbon and nutrient sequestration. Additionally, we would like to determine if there may be important opportunities for sequestration in manufactured soils with expanded applications on abandoned mine and conservation program lands.

Second, we propose the development of protocols whereby landowners could adopt organic soil management practices and quantify sequestration potential. Ultimately, this could enable landowners to participate in carbon and nutrient trading markets, which would provide a financial incentive to adopt organic soil management practices.

Third, we propose to expand the knowledge base on soil carbon sinks through communication and collaboration with other scientific, educational, research and agricultural institutions.

This is emerging as a new field from the perspective of many in the agricultural and soil management communities. While the data from the field trials is a matter of record, much needs to be done before we know how to transfer this knowledge for use in broader markets and applications. Nonetheless, what has been demonstrated is significant and shows promise in helping to reduce the build-up of greenhouse gases while promoting greater use of organic agriculture.


Resources

Bolin, B., E. Degens, S. Kempe, and P. Ketner. 1979. The Global Carbon Cycle. Wiley, New York.

Chen, Y., and Y. Avimelech. 1986. The Role of Organic Matter in Modern Agriculture. Martinus Nijhoff Publishing, The Hague.

Douds, David D. Jr, R. R. Janke, and S. E. Peters. 1993. VAM fungus spore populations and colonization of roots of maize and soybean under conventional and low input sustainable agriculture. Agriculture, Ecosystems, and Environment 43: 325-335.

Douds, David D. Jr., and P. D. Millner. 1999. Biodiversity of arbuscular mycorrhizal fungi in agroecosystems. Agriculture, Ecosystems, and Environment 74:77-93.

Drinkwater, L., P. Wagoner, and M. Sarrantonio. 1998. Legume-based cropping systems have reduced carbon and nitrogen losses. Nature 396:262-265.

Nebel, Bernard J., and Richard T. Wright. 1996. Chapter 16. Major Climatic Changes in The Way The World Works Environmental Science Fifth Edition. Prentice Hall, Upper Saddle Rive, New Jersey. 687 p.

Paul, E. A., and F. E. Clark.1989. Chapter 6 Carbon cycling and soil organic matter in Soil Microbiology and Biochemistry. Academic Press, New York. 271 p.

Puget, P., and L. Drinkwater. 2001. Short term dynamics of root and shoot-derived carbon for a leguminous green manure. Soil Sci. Soc. Am. J. 65:771-779.

Rillig, M., and S. F. Wright. 2002. The role of arbuscular mycorrhizal fungi and glomalin in soil aggregation. Plant and Soil 234:325-333.

Rillig, M., S. F. Wright, K. Nichols, W. Schen, and M. Torn. 2001. Large contribution of arbuscular mycorrhizal fungi to carbon pools in tropical forest soils. Plant and Soil 233:167-177.

Sanchez, P., M. P. Gichuru, and L. B. Katz. 1982. Organic matter in major soils of the tropical and temperate regions. Proc. Int. Soc. Soil Sci. Cong. 1:99-114.

Sedjo, Roger A. Brent Sohngen and Pamela Jagger. 1998. RFF Climate Issue Brief #12

Stevenson, F. 1982. Humus Chemistry: Genesis, Composition, and Reactions. Wiley Interscience, New York. 583.

Stevenson, F. 1985. Cycles of Soil Carbon, Nitrogen, Phosphorus, Sulfur and Micronutrients. John Wiley and Sons, New York. 380 p.

Wander, M., S. Traina, B. Stinner, and S. Peters. 1994. Organic and conventional management effects on biologically active soil organic matter pools. Soil Sci. Soc. Am. J. 58: 1130-1139.

Wright, S. F., and R. Anderson. 2000. Aggregate stability and glomalin in alternative crop rotation for the Central Plants. Biology and Fertility of Soil 31:249-253.

Related articles:
About The Rodale Institute - www.strauscom.com/rodale-facts
October 10th press release - www.strauscom.com/rodale-release
Text of the October 10th Statement of Cooperation - www.strauscom.com/rodale-MOU
About The Rodale Institute - www.strauscom.com/rodale-background

54. Glomalin, the Unsung Hero of Carbon Storage 

Another glomalin source article in my collection to share. This one relates Kristi Nichols, Sara Wright and and E.Kudjo Dzantor, both USDA soil scientists working at ARS, quantifying glomalin rates and carbon storage. Rough numbers of 1-100 mg/g give us some theoretical numbers to look for when we begin measuring our own storage sites. I have written several scientists requesting information relating glomalin to forests. Only Sara Wright has responded, only saying she wished she could work in the redwoods.
Glomalin, the Unsung Hero of Carbon Storage
ARS News Service
Agricultural Research Service, USDA
Don Comis, (301) 504-1625, comis@ars.usda.gov
September 6, 2002

Glomalin, a recently discovered major component of soil organic matter,
stores about a third of the world's soil carbon, offsetting industrial
pollution. This is according to a recent collaborative study by scientists
with the Agricultural Research Service and the University of Maryland
(U-MD) at College Park. The study was partially funded by the U.S.
Department of Energy.

The study was done by Kristine A. Nichols, a U-MD soil science Ph.D.
candidate and technician at ARS' Sustainable Agricultural Systems
Laboratory in Beltsville, Md., along with colleagues Sara F. Wright and E.
Kudjo Dzantor. Wright, an ARS soil scientist, discovered glomalin in 1996,
and Dzantor is a U-MD soil scientist.

Glomalin is a sticky protein produced by root-dwelling fungi and sloughed
into soil as roots grow. By gluing soil particles and organic matter
together, it stabilizes soil and keeps carbon from escaping into the
atmosphere. In an earlier study, Wright found that glomalin serves as a
corrective to global warming because it increases with carbon dioxide
levels.

Nichols and colleagues detected large amounts of glomalin in soils from
four states, showing it to be a major part of organic matter. The glomalin
weighed 2 to 24 times as much as humic acid, which was previously thought
to store the most carbon. But Nichols found that humic acid only stored 8
percent of total soil carbon compared to glomalin's 27 percent.

Wright has found glomalin in soils from around the world, ranging in
weight from less than 1 milligram per gram (mg/g) of sample to more than
100 mg/g. She found the highest levels in Hawaiian and Japanese soils,
indicating that some soils might be able to store large amounts of carbon
in glomalin with a turnover rate of 7 to 42 years. She is on a team
investigating underground carbon storage in tropical forests, thought to
be major carbon reservoirs.

For more on glomalin, see the September 2002 issue of Agricultural
Research magazine, online at:
http://www.ars.usda.gov/is/AR/archive/sep02/soil0902.htm

ARS is the U.S. Department of Agriculture's chief scientific research
agency.

Sunday, July 11, 2004

53. Fungal-Mediated Plant Coexistence  

This summary of articles relating elevated CO2 to increased biodiversity from CO2 Science magazine. Elevated CO2 and local plants and fungi are often enough to give native plants competitive edges in the long run. We have now shown the need for fungi in forests, the product glomalin they produce, the need for large old trees and the carbon exudates they produce, the qualities of soil glue, examples of glomalin destruction, forest practices for maximizing carbon storage at reduced risk of fire, and the critical role of this activity in restoring severely impacted watersheds.
53. Fungal-Mediated Plant Coexistence
(www.co2science.org)
Volume 6, Number 53: 31 December 2003
In a recent review of this intriguing concept, Hart et al. (2003) say "coexistence is a biological riddle, because the tendency towards competitive exclusion should favor a monoculture." Monocultures, however, are rare in nature; and Hart et al. note that several scientists (Janos, 1980; Hetrick et al., 1989; Allen and Allen, 1990; Hetrick et al., 1994) have suggested that arbuscular mycorrhizal fungi (AMF) - a common group of symbiotic soil fungi - may be "important agents promoting plant coexistence," which concept forms the basis of their review. Hart et al. begin by differentiating between two types of studies of AMF effects on plant coexistence. Coarse-scale studies are defined as those that compare the outcome of plant competition with AMF presence or absence, which they say should be "relevant to the outcome of plant interactions mainly in early successional ecosystems," where AMF "are either absent or are in low abundance and patchily distributed." Fine-scale studies, on the other hand, are defined as those that compare the outcome of plant competition when all experimental treatments contain AMF and the manipulations "involve the composition and diversity of AMF, and the ways in which they interact with plants and their soil environment." These experiments, they say, "are more relevant to later-successional situations, in which AMF are more abundant and less patchy, and the roots of most plants come into contact with them."Concentrating primarily on the latter category of experiments, Hart et al. note that "higher AMF diversity could lead to higher plant coexistence simply by increasing the probability of individual plant species associating with a compatible and effective AMF partner." In addition, they point out that shared mycelial networks "might promote plant species coexistence by equalizing the distribution of soil resources among competitively dominant and subdominant host species," noting that "soil nutrients and plant-derived carbon might flow through the network from dominant to subordinate host plants, because of a concentration gradient created initially when the dominant plant takes up more nutrients than does a subordinate plant." The former of these phenomena - the plant-to-plant transfer of nutrients - has been observed by Malcova et al. (1999) under laboratory conditions and by Walter et al. (1996) in the field. Likewise, the second phenomenon - the plant-to-plant transfer of carbon - has been observed by Grime et al. (1987), Graves et al. (1997) and Robinson and Fitter (1999).In concluding their review, Hart et al. say "our understanding of fine-scale factors is just starting to develop," and in this regard we note that many experiments conducted in recent years have added elevated levels of atmospheric CO2 to the mix of experimental variables considered within this context.Several such studies are described in our Subject Index under the heading Biodiversity (Fungi), where it may be seen, as noted in the review of Hart et al., that (1) the presence of soil fungi helps to maintain, and sometimes even increase, the biodiversity of various ecosystems, and (2) elevated levels of atmospheric CO2 help these fungi to better perform this important function. Also, under the Subject Index heading of Fungi, one can read how these principles operate in Grasses, Herbaceous Plants and Woody Plants, and how they help to enhance Carbon Sequestration.It is also interesting, in this regard, to go back to the book of Idso (1989) -- Carbon Dioxide and Global Change: Earth in Transition -- and read what he wrote about the subject nearly a decade and a half ago:"Considerable evidence may be mustered to support [an] optimistic view of the effects of atmospheric CO2 enrichment on species diversity. Looking to the past, for example, several studies of Tertiary floras have demonstrated that many montane taxa of that period regularly grew among mixed conifers and broadleaf shclerophylls (Axelrod, 1944a, 1944b, 1956, 1976, 1987), whereas today these forest zones are separated from each other by fully 1,000 m in elevation and 10-20 km or more in distance (Axelrod, 1988). Indeed, during this many-million-year period, when the CO2 content of the atmosphere was generally much greater than it is today (Volk, 1987), all three forest zones merged to form a 'super' ecosystem, which, in the words of Axelrod (1988), "was much richer than any that exists today." Even under current conditions, in fact, modern forestry experiments have demonstrated that trees planted in mixtures sometimes grow better than they do in single-species plantings (Brown and Harrison, 1983; Carlyle and Malcolm, 1986a; Carlyle and Malcolm, 1986b; Chapman et al., 1988)."One mechanism by which this type of mutualism may be fostered has to do with the stimulation of vesicular-arbuscular mycorrhizal fungi, which are ubiquitous in most terrestrial ecosystems (Gerdemann, 1968; Read et al., 1976) and the most prevalent of all soil fungi (Gerdemann and Nicolson, 1963). These unseen inhabitants of the soil provide a number of benefits to the plants they 'service.' They increase the absorption of water and nutrients by the plant, protect the plant from soil-borne diseases, and reduce the incidence of nematode infection of roots (Ingham, 1988). And as Johnson and McGraw (1988) have noted, 'their vigor may be expected to reflect the vigor of their hosts,' which with CO2 enrichment would be expected to increase. Consequently, whereas community ecology paradigms of the past, based largely on data pertaining to above-ground interactions, have tended to stress relatively short food chains with competition and antagonism as major organizing forces in community development, Edwards and Stinner (1988) note that, today, 'ecologists studying biotic interactions in soil systems generally have observed complex food webs, a great diversity of organisms, and a wide range of symbiotic interactions.' Indeed, many endomycorrhizae (Chiariello et al., 1982), as well as certain ectomycorrhizae (Reid and Woods, 1969; Read et al., 1985), have even been demonstrated to actively transfer nutrients between individual plants of both the same and different species. In fact, seedlings of some plants will not grow at all unless they interact with the mycorrhizae of an adjacent host plant (Warcup, 1988), while in other situations, both seed germination and initial plant growth rates are greatly stimulated by the presence of such fungi (Clements and Ellyyard, 1979; Masuhara and Katsuya, 1989). As a result, Moore (1988) contends that mutualism is common below ground and that it 'can have profound effects on the structure and activity of soil microbial communities, the decomposition of organic matter, and ultimately plant growth'."Yes, as the saying goes, "everything old is new again." It's been shown over and over that arbuscular mycorrhizal fungi and atmospheric CO2 enrichment make a truly dynamic duo when it comes to getting plants to "cooperate" in both maintaining and enhancing ecosystem biodiversity.Sherwood, Keith and Craig Idso
ReferencesAllen, E.B. and Allen, M.F. 1990. The mediation of competition by mycorrhizae in successional and patchy environments. In: Grace, J.B. and Tilman, D. (Eds.). Perspectives in Plant Competition. Academic Press, New York, NY, pp. 367-389. Axelrod, D.I. 1944a. The Oakdale flora (California). Carnegie Institute of Washington Publication 553:147-166.Axelrod, D.I. 1944b. The Sonoma flora (California). Carnegie Institute of Washington Publication 553: 167-200.Axelrod, D.I. 1956. Mio-Pliocene floras from west-central Nevada. University of California Publications in Geological Science 33: 1-316.Axelrod, D.I. 1976. Evolution of the Santa Lucia fir (Abies bracteata) ecosystem. Annals of the Missouri Botanical Garden 63: 24-41.Axelrod, D.I. 1987. The Late Oligocene Creed flora, Colorado. University of California Publications in Geological Science 130: 1-235.Axelrod, D.I. 1988. An interpretation of high montane conifers in western Tertiary floras. Paleobiology 14: 301-306.Brown, A.F.H. and Harrison, A.F. 1983. Effects of tree mixtures on earthworm populations and nitrogen and phosphorus status in Norway Spruce (Picea abies) stands. In: Lebrum, P.H., Andrea, H.M., De Medts, A., Gregoire-Wibo, C. and Wauthy, G. (Eds.). New Trends in Soil Biology. Proceedings of the VIII International Colloquium on Soil Zoology. Louvain-la-Neure, Belgium, pp. 101-108.Carlyle, J.C. and Malcolm, D.C. 1986a. Nitrogen availability beneath pure spruce and mixed larch + spruce stands growing on a deep peat. I. Net mineralization measured by field and laboratory incubations. Plant and Soil 93: 95-113.Carlyle, J.C. and Malcolm, D.C. 1986b. Nitrogen availability beneath pure spruce and mixed larch + spruce stands growing on a deep peat. II. A comparison of N availability as measured by plant uptake and long-term laboratory incubations. Plant and Soil 93: 115-122.Chapman, K., Whittaker, J.B. and Heal, O.W. 1988. Metabolic and faunal activity in litters of tree mixtures compared with pure stands. Agriculture, Ecosystems and Environment 24: 33-40.Chiariello, N., Hickman, J.C. and Mooney, H.A. 1982. Endomycorrhizal role for interspecific transfer of phosphorus in a community of annual plants. Science 217: 941-943.Clements, M.A. and Ellyyard, R.K. 1979. The symbiotic germination of Australian terrestrial orchids. American Orchid Society Bulletin 48: 810-816.Edwards, C.A. and Stinner, B.R. 1988. Interactions between soil-inhabiting invertebrates and microorganisms in relation to plant growth and ecosystem processes: An introduction. Agriculture, Ecosystems and Environment 24: 1-3.Gerdemann, J.W. 1968. Vesicular-arbuscular mycorrhizae and plant growth. Annual Review of Phytopathology 6: 397-418.Gerdemann, J.W. and Nicolson, T.H. 1963. Spores of mycorrhizal Endogone species extracted from soil by wet sieving and decanting. Transactions of the British Mycological Society 46: 235-244.Graves, J.D. et al. 1997. Intraspecific transfer of carbon between plants linked by a common mycorrhizal network. Plant and Soil 192: 153-159.Grime, J.P. et al. 1987. Floristic diversity in a model system using experimental microcosms. Nature 328: 420-422.Hart, M.M., Reader, R.J. and Klironomos, J.N. 2003. Plant coexistence mediated by arbuscular mycorrhizal fungi. TRENDS in Ecology and Evolution 18: 418-423.Hetrick, B.A.D. et al. 1989. Relationship between mycorrhizal dependence and competitive ability of two tall grass prairie grasses. Canadian Journal of Botany 67: 2608-2615.Hetrick, B.A.D. et al. 1994. Effects of mycorrhizae, phosphorus availability, and plant density on yield relationships among competing tall grass prairie grasses. Canadian Journal of Botany 72: 168-176.Idso, S.B. 1989. Carbon Dioxide and Global Change: Earth in Transition. IBR Press, Tempe, AZ.Ingham, R.E. 1988. Interactions between nematodes and vesicular-arbuscular mycorrhizae. Agriculture, Ecosystems and Environment 24: 169-182.Janos, D.P. 1980. Mycorrhizae influence tropical succession. Biotropica 12: 56-64.Johnson, N.C. and McGraw, A.-C. 1988. Vesicular-arbuscular mycorrhizae in taconite tailings. II. Effects of reclamation practices. Agriculture, Ecosystems and Environment 21: 143-152.Malcova, R. et al. 1999. Influence of arbuscular mycorrhizal fungi and simulated acid rain on the growth and coexistence of the grasses Calamagrostis villosa and Deschampsia flexuosa. Plant and Soil 207: 45-57.Masuhara, G. and Katsuya, K. 1989. Effects of mycorrhizal fungi on seed germination and early growth of three Japanese terrestrial orchids. Scientia Horticulturae 37: 331-337.Moore, J.C. 1988. The influence of microarthropods on symbiotic and non-symbiotic mutualism in detrital-based below-ground food webs. Agriculture, Ecosystems and Environment 24: 147-159.Read, D.J., Francis, R. and Finlay, R.D. 1985. Mycorrhizal mycelia and nutrient cycling in plant communities. In: Fitter, A.H., Atkinson, D., Read, D.J. and Usher, M.B. (Eds.). British Ecological Society Special Publication 4: Ecological Interactions in Soil, pp. 193-217.Read, D.J., Koucheki, H.K. and Hodgson, T. 1976. Vesicular-arbuscular mycorrhizae in natural ecosystems. I. The occurrence of infection. New Phytologist 77: 641-653.Reid, C.P.P. and Woods, F.W. 1969. Translocation of 14C-labelled compounds in mycorrhizae and its implications in interplant nutrient cycling. Ecology 50: 179-187.Robinson, D. and Fitter, A. 1999. The magnitude and control of carbon transfer between plants linked by a common mycorrhizal network. Journal of Experimental Botany 50: 9-13.Volk, T. 1987. Feedbacks between weathering and atmospheric CO2 over the last 100 million years. American Journal of Science 287: 763-779.Walter, L.E.F. et al. 1996. Interspecific nutrient transfer in a tall grass prairie plant community. American Journal of Botany 83: 180-184.Warcup, J.H. 1988. Mycorrhizal associations and seedling development in Australian Lobelioideae (Campanulaceae). Australian Journal of Botany 36: 461-472.

31 December 2003
Copyright © 2004. Center for the Study of Carbon Dioxide and Global Change (www.co2science.org).

52. Old Growth Needed for Sequestration  

52. Old Growth Needed for Sequestration Another article showing the slowly emerging realization of forest system mechanics. The original article appeared in the NY Times, and the response I saved from a newsgroup posting. They, again, haven’t made the glomalin connection yet. But if ever a reason was needed for saving big trees, here you go. Also, they are unaware of CO2 loss from soil disturbance.
> Do we see the writing on the wall yet ??? -Sam J.
> New York Times
> September 22, 2000
> Planting New Forests Can't Match Saving Old Ones in Cutting Greenhouse Gases, Study Finds> By ANDREW C. REVKIN
> A new study has cast doubts on an important element of a proposed treaty to fight global warming: the planting of new forests in an effort to sop up carbon dioxide, a heat-trapping gas.
> The research concludes that old, wild forests are far better than plantations of young trees at ridding the air of carbon dioxide, which is released when coal, oil and other fossil fuels are burned.
> The United States and other countries with large land masses want to use forest plantations to meet the goals of the proposed treaty. The study's authors say that any treaty also needs to protect old forests and that, so far there is no sign that such protections are being considered.
> Without such protections, the scientists conclude, some countries could be tempted to cut down old forests now and then plant new trees on the deforested land later, getting credit for reducing carbon dioxide when they have actually made matters worse.
> The analysis, published in the journal Science today, was done by Dr. Ernst-Detlef Schulze, the director of the Max Planck Institute for Biogeochemistry in Jena, Germany, and two other scientists at the institute.
> Several climate and forestry experts familiar with the work said the study provided an important new argument for protecting old-growth woods. And they say the study provides a reminder that the main goal should be to reduce carbon dioxide emissions at the source, smokestacks and tailpipes.
> In old forests, huge amounts of carbon taken from the air are locked away not only in the tree trunks and branches, but also deep in the soil, where the carbon can stay for many centuries, said Kevin R. Gurney, a research scientist at Colorado State University. When such a forest is cut, he said, almost all of that stored carbon is eventually returned to the air in the form of carbon dioxide.
> "It took a huge amount of time to get that carbon sequestered in those soils," he said, "so if you release it, even if you plant again, it'll take equally long to get it back."
> Negotiators are to meet in November to settle on methods for staving off a predicted warming that could disrupt ecosystems, harm agriculture and cause sea levels to rise, eroding coasts.
> The negotiations are taking place under the Kyoto Protocol, an agreement that was signed by more than 100 countries in 1997 but has not yet been ratified. It sets goals for cutting greenhouse gas emissions starting in 2008 but includes few details on how to achieve them. The United States, Canada, Russia and other countries have been pressing to achieve as much as half their greenhouse gas reductions not at the source but by using "sinks" like forests to remove carbon dioxide.
> In the last round of talks, which ended last week in Lyon, France, some countries were still seeking treaty language that could allow some new planting to occur on land that was recently cleared of old forest and get credit for greenhouse-gas reductions, said Mr. Gurney, who attended the talks as an observer.
> David B. Sandalow, an assistant secretary of state who was the chief American delegate in Lyon, said that the treaty drafts so far could theoretically allow such a practice but that the United States was seeking to prevent this.
> "We're committed to protecting old growth and finding ways to address this issue," Mr. Sandalow said. The German study, together with other similar research, has produced a picture of mature forests that differs sharply from long-held notions in forestry, Dr. Schulze said. He said aging forests were long perceived to be in a state of decay that releases as much carbon dioxide as it captures.
> But it turns out that the soils in undisturbed tropical rain forests, Siberian woods and some German national parks contain enormous amounts of carbon derived from fallen leaves, twigs and buried roots that can bind to soil particles and remain for 1,000 years or more. When such forests are cut, the trees' roots decay and soil is disrupted, releasing the carbon dioxide.
>Centuries would have to pass until newly planted trees built up such a reservoir underground. New forests are fine as long as they are planted on land that was previously vacant, Dr. Schulze said, adding, "but there has to be a focus on preserving the old growth."
Copyright 2000 The New York Times Company
>
While the article is correct, it is somewhat misleading. The carbon locked up underneath the soil is actually *given* by the tree to the mycorrhizal fungi associated with the tree. This carbon is invested in
producing fungal fruiting structures and (in some cases) increasing the volume of soil where these fungi are located. Fungi can extend literally hundreds, if not thousands of feet away from the host tree roots. They
are fine enough to penetrate tiny cracks and crevices in rock, and may extend downward to the water table, thus becoming an important adjunct to the long-term survival of a tree.
However, these fungi often have considerable die-out each year. Fortunately, the fungi produce such prodigious quantities of spores (an"average sized" Rhizopogon may contain 20 million spores, for example)
that over time the rhizosphere expands a little each year.
Much of the effective bulk of carbon considered to be a carbon sink is stored within the tree cell walls itself. Recent research by Chris Maser, J. Trappe and others suggests that a 4-foot diameter Douglas fir of approximately 300-years-age would take at least that long to degrade. Thus, for every 1 year of increased age, it walls away that much more carbon.
Wood degredation is typically by fungi also, but includes soil organisms, fungi, and insects in a complex ecosystem itself.
It is also true that cutting larger trees into lumber for long-term house construction material similarly walls-away carbon from the environment, provided that the home is well-constructed and designed to last for at least 80 years. The problem with home construction is that it takes larger-diameter wood (4' diameter) and converts it into smaller-diameter wood (2x4's), which then tend to degrade much faster than the original larger sized wood.
So you see, both sizes of trees are important, both as carbon-sinks for the future and as timber for today. The public _must_ be aware, however, that trees are growing organisms. Eventually they die. And somewhere between clearcutting and no cutting there is probably a nice annual harvest rate which does not seriously impact either forest health or forest ecology.
Of course, if foresters aren't growing fungi it's hard to see how they can claim to be growing trees, since mycorrhizal fungi are so important to tree health and growth.

Daniel B. Wheeler
www.oregonwhitetruffles.com

Sent via Deja.com http://www.deja.com/

51. Growth Rates and Growth Periodicity of Tree Roots 

51. Growth Rates and Growth Periodicity of Tree Roots
Another fine article on the Kelsick website, this time about roots. They ignore mycorhizzhia while telling us they are ignoring it. This is why we have to piece this case together with articles based on solid references. http://www.chesco.com/~treeman/.html
Growth Rates and Growth Periodicity of Tree Roots [International Review of FORESTRY RESEARCH vol 2 page181-page 236 1967]
HORST LYR AND GUNTER HOFFMANN
Institut fur Forstwissenschaften, Eberswalde, der Deutschen Akademie der Landwirtschaftswissenschaften zu Berlin (DDR)

I. Introduction
For a long time the root system was regarded only as an auxiliary organ of the plant providing mechanical fastening in the soil and absorbing water and mineral salts. These functions are indeed important, but it should be kept in mind that roots are highly specialized organs in which numerous syntheses are performed (Mothes, 1956). Water uptake and mineral absorption, according to modern concepts, are closely related to metabolic activity and growth of roots. From many practical experiences it is well known that vigorous root growth is necessary for good shoot growth and that disturbances in root growth impair shoot growth.
In spite of this, relatively little is known about growth behavior of roots. This is mainly because of difficulties arising from the methods used. Many failures in cultural techniques result from ignorance of the normal course of root growth and of the influences of environmental conditions upon it. Numerous questions are still almost uninvestigated,-for example, the correlative reciprocal effects between root and shoot systems, the influence of environmental factors on the intensity and course of root growth, and specific optimum values for the different tree species.
At present it is difficult to draw general conclusions from the statements in the literature because environmental conditions and research techniques have varied widely and often have been only inadequately described. As is described in detail in the last part of this article, practical conclusions could be drawn from better knowledge about root growth. The possibilities in this field will be much expanded with increasing knowledge on the specific requirements of various tree species for good root growth. Optimal conditions for vigorous root growth are as important for high productivity as are good conditions for shoot growth. Although this has often been said, little is really known in detail. The physiology and ecology of root growth are still a neglected segment of plant physiology.

II. The Root System and Growth Rates of Tree Roots
A. The Root System

1. THE GROSS ROOT SYSTEM
Many investigations on the structure of the root system of trees have been performed during recent decades to get information on the specific peculiarities of root morphology and on the extent of utilization of the soil volume by trees growing on different sites. Valuable knowledge was obtained by such static surveys on trees of different ages and on different sites. By investigating root arrangement and distribution in the soil, practical conclusions were drawn for silviculture and horticulture. Only some of the most important papers on this subject can be cited here.
The root system of Fagus silvatica was studied by Zielaskowski ( 1898), Krauss and co-workers (1934, 1935, 1939), Bonnemann (1939), Krahl-Urban (1951), Petsch (1955), and Hausdorfer (1959). That of Pinus silvestris was investigated by Tolsky (1904), Aaltonen (1920), Kokkonen (1923), Liese (1926), Hilf (1927), Laitakari (1929), Wagenhoff (1938),Simanjuk (1950), Kalela (1950), Rachtejenko (1952), Yeatman (1955), and Hausdorfer (1959). Concerning the root system of Pseudotsuga taxifolia Britt. there are the papers of Groth (1927), Wagenknecht (1958), and McMinn (1963); for Picea abies there are the reports of Vater (1927), Wagenknecht and Belitz (1959), Wiedemann (1927), Krauss and co-workers (1934, 1935, 1939), Kern et al. ( 1961 ), and Melzer (1962b). The root system of Quercus borealis var. maxima was studied by Lemke (1955). In addition, information on several other tree species, mostly North American, may be found in the papers of Holch ( 1931 ), Biswell (1934), Coile (1937), Scully (1942), Joachim (1953), Kreutzer (1961), Dchjen (1962), and Lyford and Wilson (1964). Kreutzer (1961) and Heikurainen (1964) have reported on root development of some tree species under the influence of soil water conditions which were changing as a result of amelioration measures.
The results show that it is difficult to establish general, intraspecific rules of root development because site and soil conditions modify root formation to such a degree that peculiarities of the species are partly or entirely obscured (Wittich, 1947; Wagenknecht, 1955). In spite of this limitation available knowledge of root development has allowed some conclusions to be drawn about the practical utilization of trees in silviculture and horticulture, conclusions which are of special importance for soils with different layers or with high resistance to the penetration of roots (Kvarazhelia, 1931; Wagenknecht, 1955; Kreutzer, 1961).
In many cases structure and reactivity of the root system are of decisive significance in determining site latitude and site tolerance of a tree species. The early development of a plant is in especially close correlation with its ecology, because it is often decided in the seedling stage whether a species is able to colonize a particular site. Quick reaching of deeper soil layers which are not in danger of drying out, and a sufficient ability to compete with roots of other species, are decisive factors in natural and artificial reproduction. Tree root systems consist of various types of morphologically and functionally different roots. The most generally used nomenclature for the different parts of the root system is given in a schematic way in Fig. 1.
Some tree species have a tendency to form taproots. This is most obvious in seedlings and is often found in species with large seeds rich in reserve foods, such as Quercus, Carya, Castanea, and Juglans. In these the taproots also serve as storage organs for the seedling. Taproots may sometimes be observed in other genera, notably Pinus. They are well suited for quickly reaching greater soil depths and are quickly and preponderantly developed during the juvenile phase at the expense of the storage material of the seeds, whereas the development of the shoots is at first rather slow and sometimes restricted to one or very few leaves, especially so in some desert plants (Kausch, 1959).
Taproots are able to reach low ground-water levels and secure the water supply of a tree even in dry areas. Such roots can grow down to considerable depths; for example, l8-year-old apple trees to 10 m (Wiggans, 1936); Robinia to 20 m (Schimper and Faber, 1935); Prosopis to 15 m; and Tamarix to as much as 30 m (Kausch, 1959). A strong horizontal. For diameter classes the classification of Grosskopf (1950) and Kreutzer (1961) is proposed (approximately equivalent English terms are given here): finest roots, < 0.5 mm; fine roots, 0.5 - 2 mm; weak roots, 2 - 5 mm; firm roots, 5 -10 mm; rough roots, 10 - 20 mm; strong roots, > 20 mm. According to Grosskopf (195O), "fine rot capacity" means root weight or length of roots from
0.5 to 2 mm per liter of soil; "finest root capacity" means roots <0.5 mm in diameter per liter soil.
root development may be observed on poor sites (such as sand dunes) ; where Pinus, Betula, and Robinia may form roots 10, 20, or even 40 meters long, which often results in considerable root competition. Roots, of Acer rubrum may also reach a length of 25 m (Lyford and Wilson, 1964).
A shallow, superficially expanded root system, which is typical of Picea species, is very effective in absorbing the ephemeral moisture after rains or melting snow, especially on shallow, rocky soils, which may be an ecological advantage in the mountains or in deserts. In the Cactaceae a superficial root system is combined with an astonishing ability for quick regeneration of root tips, which is a special mechanism of drought resistance (Kausch, 1955).
In orchard trees the horizontal expansion of the root system is normally one and a half to two times as large as the crown diameter (Kolesnikow, 1962b). In some cases it amounts to three to five times the crown radius ( Kvarazhelia, 1931). This seems to be true also for other trees growing as solitaires, but the relations may be strongly modified by the environment. This should be noted in practical fertilization of older trees, in which the highest concentration of absorbing fine roots occurs at some distance from the trunk. It should also be kept in mind that there exist differences in root morphology, not only between tree species, but also between their provenances, which may be correlated with differences in growth performance. This was demonstrated in Pinus silvestris by Bibelriether (1964).

2. THE FINE ROOT SYSTEM
Investigations of the root system often concern only the gross root system. Although valuable conclusions may be drawn, it should be kept in mind that the quantity and activity of fine roots are primarily decisive for water and mineral salt supply of a tree. At present only a few investigations have been published in this field (e.g. Coile, 1937; Grosskopf, 1950; Grunert, 1955; Hausdorfer, 1959; Kern et al., 1961; Buchholz and Neumann, 1964; Zottl, 1964; see the latter for further literature).
Diameter analyses of roots of young plants show that the major part of the root system consists of fine roots. From Table I it may be seen that in the four tree species investigated in detail, although only 14 to 60% of the total root weight is represented by fine roots under 1 mm in diameter, these make up 86 to 99% of the total root length. The proportion of the total root surface represented by fine roots is, of course, much higher still. As may be seen from the investigations of Coile (1937) or Hausdorfer (1959) in soil profiles, these relations are similar in mature trees, although the weight relations may be different.
Root weight is a less suitable index than root length. Length is better correlated with surface development, a most important. factor affecting physiological activity and exchange of substances (Grosskopf, 1950).
With increasing soil depth the proportion of fine roots increases. The different percentages which the single diameter classes contribute to total root weight and length in Table I show specific peculiarities in the root formation of the four tree species.
From our knowledge of root quantities, no direct conclusions about root activity can be drawn, although a loose correlation may be expected to exist. At present, detailed, comparable investigations are lacking. Surely the activity of mycorrhizae is very important. But this activity depends partly on the fungal species, which fact complicates the problem (Ritter and Lyr, 1965).
Measuring and recording the static state of the root system may be of value in solving many questions. From a physiological point of view, however, the dynamic of root development is of greater importance. Because of unequal penetration of precipitation through soil fissures, old root tubes, and differently permeable soil areas, and because of the local un-equal water uptake of roots, the soil is frequently very inhomogeneously wetted. Furthermore, the capillary water conduction of most soils is too low to equalize the differences quickly enough, so that with increasing drying out the water supply available to absorbing roots is essentially a stagnant one. New water and mineral salt sources must be found by active root growth, which under such circumstances is an important factor in ecological competition. A high rate of root increment means the penetration of a large soil volume and the accessibility of water reserves in the soil which is important for most trees in maintenance of a stable water regime. Deficiency of water and nitrogen leads to a promotion of long root growth and restriction of side-root formation. This favors a fast penetration of the soil.
Because of technical difficulties, few attempts have been made to measure root growth continuously. Therefore little is known concerning maximal and average root growth rates of different trees under similar environmental conditions. The influence of defined environmental factors on root growth is still rather obscure. The data of different authors diverge widely because of different methods and conditions.
In Table II some data from our own measurements and from the literature are summarized. It can be seen that roots of fast-growing trees may reach values which are not inferior to that for herbaceous plants. Even the average values are considerable. It must be noted that the values measured in the root laboratory were obtained from newly planted trees. The disturbed root/shoot relation is normalized during the first year by preferential root growth, and a high incremental rate is reached. The "average" value means the average growth rate from all growing roots. This is not the average of the whole root system, because not all roots are growing at the same time. The data of different authors are not strictly comparable, but the differences in methods cannot be discussed here.

4. DEPTH GROWTH
Little is known about the course of depth growth because excavations disturb root growth, and observations in Sachs-type root boxes are possible only for a short time and must be restricted to seedlings. In the root laboratory (Fig. 2) at Eberswalde*1 an adequate survey on time course, periodicity, and intensity of root growth could be obtained.

*1 Eberswalde is located at 52° 50' north latitude and 13° 49' east longitude; altitude 30 m above sea level; 572 mm annual mean precipitation (March, 38 mm; July, 81 mm); 8.4°C annual mean temperature; 18.8°C annual temperature variation. Characteristic differences in root development showed up in the first vegetative period after planting.
Because a homogeneously compacted sandy soil was used in the root laboratory, peculiarities of the tree species in depth growth were easily recognizable. In Figs. 3 to 5 the course of shoot and root growth are plotted for Betula pendula Roth., Populus euramericana "I 214," and Pinus silvestris L. Each point represents a growing root tip (see also Figs. 18 and 19 for Robinia pseudoacacia L.). It is obvious that depth development is very different in different species.
Whereas the roots of Populus and Robinia during the first vegetation period advance to a depth of 1.5 to 2 m, those of Pseudotsuga taxifolia, Pinus silvestris, and Larix leptolepis reach a depth of only 0.5 m. Quercus borealis var. maxima and Betula pendula go a bit deeper, whereas Picea abies remains shallower. Some values for maximal depth growth during the first vegetative period of comparable plants are summarized in Table
II. The rate of depth growth is partly correlated with the general growth rate of a tree species (see Populus, Fig. 4). Beyond that, specific differences exist in the rate of depth penetration. The fast depth growth of tree seedlings with taproots has already been mentioned. But some trees with typical heartroot, rather than taproot, systems for example, Betula pendula or Robinia pseudoacacia-soon reach even greater depths, which is very important for their water supply. Strong growth during youth means more than escape from shading. In some species deep rooting also diminishes danger from drought periods and so increases the ability of the species to compete. In Betula papyrifera shallow rooting on some sites leads to die back (Pommerleau and Lortie, 1962).
Figures 3 to 5 show that as growth proceeds, the center of most active root growth (statistically represented by the solid line) shifts to deeper soil layers. This corresponds to the normal tendency of expansion of the root system under the influence of a specific correlative regulation. In virgin soils deviations are to be expected because of the irregular deposition of mineral salts. Their enrichment in the upper soil 1ayers with high humus content leads to a concentration of fine roots in the upper horizons and thereby to a shallower rooting of most trees growing on such soils. Under ordinary outdoor conditions as much as 80 to 90% of the total mass of fine roots may be found in the upper soil layers (Coile, 1937; Scully, 1942; Hausd6rfer, 1959).
In newly planted trees the tendency of the root growth center to shift to greater depths is very common. In older trees, which have fewer possibilities for further expansion of their root systems, other relations are found. In these, "growth nests" are formed around some strongly growing roots. Some roots stop growing and die; others form regeneration roots so that a more inhomogeneous growth distribution results. This can sometimes already be seen in the third vegetation period after planting, as, for example, in Pinus silvestris (Fig. 6). Here root growth starts and stops irregularly at different depths, and the density of growing roots is lower than in younger trees (although the total amount may be larger). Kinman (1932) made similar observations in orchard trees.
Aside from the expansion of a tree's root system, its density is probably of great importance to the tree's ability to compete. In Figs. 3 to 5 (and in Fig. 18 and fig. 19) distinct differences in the density of growing fine roots are evident. Penetration of the soil is very intensive in Larix leptolepis, Betula pendula, and Pseudotsuga taxifolia. A weak fine root formation is characteristic of Quercus borealis var. maxima, whereas Robinia pseudoacacia and Populus euramericana have an intermediate position. (In Populus the very fine hair roots with a diameter of less than 1 mm, which are typical of this species, were not registered. If these had been included the Populus root system would have been very much enlarged. See Table I.)
A low density may be compensated for by mycorrhizae formation, which is not considered here. The fine fungal hyphae may penetrate - more or less intensively, depending on the fungal species-the between- root soil spaces and thus make accessible nearly the whole soil volume of the root zone. However, this is of importance only in the upper soil layers, because frequency of mycorrhizae decreases with increasing soil depth (Preston, 1942; Werlich and Lyr, 1957). This may not depend on the humus content and the aeration of the soil, but seems to be caused by the physiological state of the different parts of the root system. For example, the nodule formation in Robinia pseudoacacia grown in the open and in the root laboratory shows very similar relations (Hoffmann, 1960; Lyr, 1963).
6. ACTIVITY IN SOIL PENETRATlON Tree roots are forced to penetrate large soil volumes, often against the mechanical resistance of densely packed soil layers, especially when the trees are acting as pioneer plants. In normal stands, young tree roots may follow old root channels formed by rotted roots or may grow in chinks of loam or other soil crevices and fissures. From practical experience it is well known that different tree species exhibit different activities in penetrating compacted or dense soil horizons, which on certain sites is the decisive feature for the choice of a tree species.
Results gained from comparable experiments on root activity, as the ability for penetration of dense soil layers is often called, are rare. An outstanding piece of work is that of Leibundgut et al. (1963) on the penetration of seedling roots through artificially made compacted clay layers in boxes. On the basis of measurements of the percentage of the total root mass in the clay and in the loose layer beneath at the end of the vegetation period, the following sequence was determined: Quercus robur (26%), Alnus incana (8.7%), Alnus glutinosa (8.0%), Carpinus betulus (2.6%), Picea abies (1.6%), Pseudotsuga taxifolia (1.5%). This corresponds well to the ranking of root activity observed in the same species growing in the open. Species with a high root-to-shoot ratio seem to have a greater ability to penetrate hard soil layers. Gardner and Danielson (1964) in their experimental investigations found that optimal aeration of the soil and optimal water content of the root zone increased the ability of roots to penetrate.
7. LONGEVITY OF FINE ROOTS
Statements and opinions expressed in the literature on longevity of fine roots are very divergent. According to Kinman (1932) fine roots may die, although only days old, when the base root begins to form periderm. In other cases longevity was estimated to be a few weeks (Childers and White, 1942). Heikurainen (1955) suggested, on the basis of his studies, a longevity of roots with diameters under 1, 1 to 2, and 2 to 5 mm of 3, 5, and 10 years, respectively. In general, fine roots seem to live at least through one vegetative period, which, however, on no account is the maximal age under favorable circumstances. In the root laboratory at Eberswalde, mycorrhizae and fine roots 2 years old, and older, could be observed. The longevity of fine roots depends partly on the correlatively regulated distribution of assimilates and partly on the growth intensity of the roots of higher rank. In the open a large fraction of the fine roots may be killed periodically by drought or frost, especially in the upper soil layers, so that frequent regeneration may consequently occur. This being the situation, it is obvious that any generally valid statements on the longevity of fine roots are difficult to formulate.
B. Periodicity of Root Growth
In addition to growth rate, growth periodicity of tree roots is also of scientific and practical interest. Whereas detailed information, and in some instances reproducible results, have been obtained on shoot growth, reports on root growth are in disagreement or are contradictory. At present it is still difficult to discriminate between peculiarities of species and environmentally induced reactions. For centuries investigations of the course of root growth have been made from a practical viewpoint, but owing to difficulties with techniques conclusions have been very vague. Theophrastos of Lesbos (372-287 B.C.) observed that roots start growing before shoots in spring. Of the older works the following should be cited: Hales (1748), Du Hamel du Monceau (1758), von Dieskau ( 1776), G. L. Hartig (1808), Konig (1820), Lindley (1855), Th. Hartig (1863), von Mohl (1862), and Nobbe (1862). Comprehensive monographs have been compiled by Resa (1877), Wieler (1894), Engler ( 1903 ), MacDougal (1938), Ladefoged (1939), and Reed (1939).
The first contributions to our knowledge of root growth were in most cases based on casual observations during excavation or planting of trees. Systematic study of the subject was begun by Th. Hartig (1863) and Resa (1877). Hartig, specifically, gave the impulse for studies on morphological variations of the root system, and Resa for investigations on growth periodicity. Resa's method consisted in making periodic excavations of tree roots in their natural habitat to determine whether growth was or was not occurring. Similar methods were used later by Petersen (1898), Tolsky (1901), McDougall (1916), Stevens (1931), Reed (1939), Ladefoged (1939), and Vorobieva (1961).
Since the work of Stevens (1931) attempts have been made to get quantitative informations by marking the root tips. Of course only very incomplete information could be obtained by these methods because changes in soil structure and crushing and irritation of roots disturb their growth. When conclusions concerning active growth are drawn from the existence of white root tips, differences in browning of root tips introduces additional inaccuracies. Heikurainen (1955), Kalela ( 1955), and Kolesnikow (1962a ) have periodically determined root weights from soil blocks and in this way have obtained indirect data on root growth.
Busgen (1901) was the first to use root boxes for investigations on growth periodicity of forest trees. Because of size limitations only small plants can be investigated by this method. Nevertheless, it was employed later by Engler (1903), Crider (1928), Bodo (1926), Woodroof and Woodroof (1934), and L. M. Turner (1936). To permit continuous measurements of roots in their natural habitat Kinman (1932) and Rogers (1935) made trenches near orchard trees, set framed glass windows against the soil profile, and covered the pit between the measurements.
In our own investigations a root laboratory (Hoffmann 1966a) has proved to be valuable for plants from 3 to 8 years old. It consists of twelve root boxes (Fig. 2) measuring 1 X 1 m, with a depth of 2.2 m. The construction of some of the root boxes allows the introduction of measurement instruments, application of chemicals, and taking samples of roots or soils. Roots with diameters of 0.5 to 1.0 mm are measured at intervals of 1 to 2 days. Simultaneous determinations of the height growth of the main shoots are made. According to the size of the experimental plants, each root box contains 9 to 15 trees. Of course, only a part of the root system is visible through the glass panels; therefore measurements have in part been supplemented by determination of total amounts of root masses at the end of the vegetation period. Because of temperature differences and some influence of light, root growth in the interior of the box may be a bit different from the visible root growth. This had already been recognized by Engler (1903), but in the root laboratory these effects are of little influence. When trenches are used in the open, the regeneration arising from injured roots often gives a false picture of normal root growth (Kinman, 1932).
In all root growth measurements "longroots" have been used exclusively. These are the "Langwurzeln" of Busgen (1901). Such roots have also been called "Triebwurzeln" (Bodo, 1926), "main roots" (Rogers, 1935), "growth roots" (Kolesnikow, 1962b), and "rope-like laterals" (McQuilkin, 1935). Growth measurements of roots of higher rank are extremely difficult because of their large number, their limited growth, and their small size. Here only determination of total weight gives reliable values.
1. GROWTH INITIATION
In general, roots of trees in temperate latitudes have a period of rest in winter (this will be discussed in detail in the next section). Growth is resumed in spring. The starting time depends on the tree species and the weather. At present an exact theoretical base for a prognosis of the initiation of root growth from climatic data is still lacking. Richardson (1958), however, made a fairly successful attempt with Acer saccharinum.
For a general theoretical elucidation of root rest and growth resumption more experimental data are necessary. Probably it is a complex process, in which hormonal relations between root and shoot are very important, but hormonal regulation of root growth is in many aspects still obscure (Torrey, 1956; Romberger, 1963). Furthermore, knowledge of endogenously and exogenously influenced periods of dormancy of the individual organs of trees is limited.
Most authors agree that root growth starts before shoot growth, which was observed by Theophrastus 2250 years ago. Time between root growth initiation and expansion of swelling buds is extremely variable and depends on environmental influences and on specific physiological optima. As detailed observations show, and the experiments of Richardson (1958) confirm, an impulse from the buds (which in this phase show a slight swelling) is necessary for root growth initiation. Apparently auxins are transported from the shoot to the root, which starts growing earlier because of a lower temperature optimum. According to Richardson (1958), roots of Acer saccharinum resume growth at 5°C, whereas bud expansion begins at 10°C. Similar differences probably exist in other tree species, but the absolute values may be expected to be different from species to species, and even from provenance to provenance. By February (in Central Europe) most trees have overcome endogenous dormancy and have entered postdormancy or the state of readiness (quiescence) in which temperature determines the time of bud opening. Increased soil temperatures lead directly or indirectly to earlier root growth, whereas bud expansion is not influenced (see Section II,C,l,a and Fig. 10).
The genus Larix is an exception to this general behavior, as was earlier demonstrated by the data of Engler (1903) and has been confirmed by our own measurements. In Larix species, needles of short shoots are unfolded long before root growth begins. These new short-shoot needles probably have the function of synthesizing the necessary assimilates for root and shoot growth, because reserve food storage in Larix is rather limited. Long-shoot expansion begins some time after needle unfolding and root growth initiation (Hoffmann, 1966a).
The beginning of root growth in Quercus borealis var. maxima is also relatively late. In Europe it may coincide with leaf unfolding. In most trees the first root growth is made at the expense of reserve materials. Therefore differences may exist between young seedlings and older trees in the continuation of root growth during shoot dormancy. Evergreen conifers, which can photosynthesize during winters, seem to behave differently from deciduous trees. In Pinus silvestris, root growth continues or is resumed independently of shoot growth when soil temperature is high enough (unpublished observations).
2. GROWTH PERIODICITY DURING THE VEGETATION PERIOD
Opinions on root growth rhythm during the vegetation period are very divergent. This is not surprising, if one takes into consideration. the fact that the results were obtained from different tree species in different areas and climates and with different methods. Instead of the theoretical "normal distribution" or bell curve of mass increment, root growth as well as shoot growth has a very irregular time course. Resa (1877) assumed an antagonistic interrelation between root growth and shoot growth. This means that in months with strong shoot growth only a limited root growth or none at all should take place, and vice versa. In contradiction to this theory, Wieler (1894) and Busgen (1901) pointed to a functional relation between root and shoot and argued that strong root growth must be bound to strong shoot growth. On the basis of comprehensive measurements Engler (1903) came to the opinion that in Central Europe all tree species have a maximum period of root growth in May and June, which is interrupted in August by a rest period, followed by a second but lower peak in autumn. Such a diminution or interruption of root growth in midsummer has often been described (McDougall, 1916; L. M. Turner, 1936; Kolesnikow, 1962b). But other authors (Tolsky, 1901; Hesselink, 1926; Rogers, 1935; Roze, 1937; Reed, 1939; Ladefoged, 1939) found no typical growth curves with two distinct peaks, which agrees with our own investigations.
We regard midsummer root growth cessation as due to unfavorable environmental conditions (periods of drought or high temperature). Under equivalent environmental conditions the rhythm of root and shoot growth differs from species to species and even from one individual tree to another. Some comparable curves for several tree species are summarized in Fig. 8. Because of the changing weather conditions, growth curves for different years are quite divergent, and generalizations are not yet possible. In our own measurements, neither significant antagonistic nor synergistic interrelations between shoot and root growth could be found by mathematical analysis. Interpretation is further complicated by the fact that at the same time growing and nongrowing roots may be found (Stevens, 1931; Ladefoged, 1939; Wilcox, 1954), so that their proportion should be determined. The correlation between environmental conditions and root growth is probably very complex, because besides direct influences of soil factors many indirect influences may act upon roots via primary effects upon activity.
Only the most general rules of root growth in a moderate climate can be stated here. Maximal root growth, with regard to both the number of growing roots and the total growth in length, in most tree species occurs in the early summer (June and July) (Fig. 8). Seedlings with early termination of shoot growth (Quercus type) often exhibited strong root growth in midsummer. Regarding the growth rate of individual roots, a maximum in early summer is evident, especially in deciduous trees (Populus, Robinia, Quercus, etc.), whereas conifers (Pinus silvestris, Picea abies, Larix decidua, and Pseudotsuga taxifolia) show a more uniform growth throughout the whole vegetation period, (Fig. 8). During August, but especially in September, root growth begins to diminish (Hoffmann, 1966a). Shoot growth in most species has stopped by the end of August or early September, often much earlier. Towards the end of the vegetation period the number of growing roots decreases considerably, and longer pauses in growth of individual roots can be observed. The measured values are often represented by only a very few roots.
Some species show peculiarities in root growth. For example, in Pinus silvestris root growth is very weak during the time of formation of new shoots and needles and increases considerably after expansion of the needles. This may be true also in other Pinus species; it seems to be correlated with the strong consumption of assimilates by the growing shoot. During this growth phase negative balances of assimilation have been measured (Neuwirth, 1959). .
3. TERMINATION OF ROOT GROWTH
In areas having low winter temperatures, root growth usually stops in the autumn. In the climates of Eberswalde, root growth of most trees ceases in September or October; only in some years does growth continue into November or even December. This was observed in 1963 in Betula pendula and Pinus silvestris. In all species root growth continues longer than shoot growth and can go on after leaf abscission. This is remarkable, because at this time (end of September or beginning of October) shoots are often already in deep dormancy (Vogl and Kemmer, 1961). Evidently there exists a certain autonomy of root growth, and it may be that- contrary to the condition in shoots-no internally controlled period of dormancy is present in roots. This is confirmed by observations that artificial heating of the soil extends root growth considerably (see Fig. 10) and that keeping of trees (Pinus strobus) in a warm greenhouse leads to continuous root growth (Stevens, 1931).
Several authors have described a winter growth of roots. This seems to be restricted to regions with mild winter temperatures and frost-free soils. It was observed for conifers and deciduous trees in the southern part of the United States, in British Columbia, in the Crimea, and in parts of Europe (Du Hamel du Monceau, 1758; Harris, 1926; Crider, 1928; L. M. Turner, 1936; Kolesnikow, 1962b). Apparently both evergreen trees and deciduous trees may have an uninterrupted root growth under certain circumstances. But in all cases a diminution of growth rate and the number of growing roots during the winter period has been reported.
4. DIURNAL GROWTH RHYTHM
Very few data exist on diurnal growth rhythms of roots. Kolesnikow ( 1962a) mentioned that growth should be stronger at night than during the day. Exact measurements in the root laboratory at Eberswalde have confirmed these statements. Although active root growth occurs during both day and night, it is more rapid at night. With day growth of a species taken as 100%, night growth of the same species gave on an average the following relative values: Populus trichocarpa 160; Quercus borealis var. maxima 137; Pinus silvestris 136; Picea abies 130. The diurnal rhythm of shoot growth is much more variable. Whether diurnal growth rhythms are caused by internal periodicity or only by the externally regulated periodicity of photosynthesis, translocation, and transpiration has not yet been investigated.
C. Root Growth and Environmental Conditions
1. ROOT GROWTH AND SOIL TEMPERATURE
Because root growth, as well as mineral salt and water uptake, is dependent on metabolic processes, it is to be expected that soil temperature will influence root growth and activity and indirectly whole plant growth also. Shoots are dependent on roots for a sufficient supply of water, minerals, and some organic compounds. According to Tew et al. (1963), soil temperature has an even greater influence on transpiration than have air temperature and humidity.
Investigations on the effect of soil temperature on root growth are complicated by the fact that growth intensity of roots depends not only on temperature-as it might in heterotrophic microorganisms on an optimal medium, or in isolated roots in artificial culture-but also on soil moisture and shoot activity (carbohydrate supply), which are themselves influenced by light, air temperature and humidity, and root activity. Therefore no simple dependence of root growth rate on soil temperature can be expected under natural conditions. As our own measurements show, the rate of root growth during the vegetation period changes much more than soil temperature. The latter is a dominant factor only in spring and fall, because it acts together with other factors, such as soil moisture and shoot activity, during the summer.
Most authors agree that optimal temperatures for roots are lower than for shoots of the same species. This is demonstrated by the experiments of Richardson (1958), for example.
a. Cardinal Temperature Values. It is difficult to give useful values for minimum, optimum, and maximum temperatures for root growth of trees. Most authors have not distinguished between a physiological and an ecological optimum and have neglected the influence of other factors on these cardinal values. The method of measuring growth is very important in determining the temperature values. Therefore most data are not strictly comparable. Short-term measurements of root growth rates (Ladefoged, 1939) reveal the physiological optimum temperature (Fig. 9), which can also be obtained with isolated cultured roots. Ecological optimal values, on the other hand, are dependent on the carbohydrate balance and other factors. A long period of higher soil temperature can lead to a negative carbohydrate balance because of enhanced root respiration. In many cases the ecological optimum, therefore, lies below the physiological optimum. In Table III some cardinal temperature values from the literature are summarized. It can be seen from these data that the range of temperature in which growth is possible lies between +2° and +35°C.
Distinct differences in cardinal temperature values exist between species. This is probably true for provenances also. According to Aaltonen (1942), Pinus silvestris is more thermophilic in its root growth than is Picea abies.
The most exact data are those concerning the minimum temperature, because its determination from growth initiation or growth cessation is relatively simple. But even here the physiological and ecological values may be different. Picea abies, Abies alba, Fagus silvatica, and Acer pseudoplatanus evidently have a rather low minimum. Growth begins or stops in the vicinity of 2° to 4°C. On the other hand, Citrus species begin root growth only when the temperature rises above 11°C (Muromtsew, 1962). Roots of some Malus varieties and Prunus persica may even be called thermophilic.
A comparison of optimum temperatures from the literature is at present nearly impossible because physiological and ecological values are confused and the definition and determination of an ecological optimum are still obscure. Physiological data give information on the temperature at which the highest growth rate of roots has been observed, but it does not follow that heating the soil to such a temperature would give an optimal effect on an ecological time scale. Here further investigations are needed.
It is striking that most physiologically optimal values lie above 20°C, temperatures which only rarely are reached in the soil. Although the ecological optimum may be lower than the physiological, it must be expected that on many sites soil temperature is suboptimal. In some cases it can even be the limiting factor. In our own experiments with Robinia pseudoacacia, artificial soil heating (increase of temperature about 5°C above normal) induced root growth which began 41 days earlier and terminated 44 days later than in control plants, whereas shoot growth (normal climate in the open) began only one day earlier (Fig. 10). The growth rhythm of the shoots in the two groups was similar, but the final height of shoots with soil heating was increased by 15% (11 cm). This effect seems to have been caused by the enhanced metabolic activity of the roots. Because R. pseudoacacia is a thermophilic tree species, further experiments are needed to show whether this is true for other species also.
In nurseries, soil mulching with plastic film can be used to increase soil temperature and conserve moisture. Sorgum vulgare showed an in- crease In yield of 68 to 76% after such treatment (Pusztai, 1963). However, no such data for tree species are yet available.
The effect of temperature on root growth is complicated by interrelations of root and shoot temperatures. Low root temperatures and high shoot temperatures favor shoot growth, and the inverse also holds, but high root or shoot temperatures can counterbalance low root or shoot temperatures to some extent (Hellmers, 1963).
The maximum temperatures are of practical importance only in special cases. In hot and dry regions (semidesert afforestations) root growth may be limited by high temperatures. This may also happen in temperate latitudes during dry periods in the season of strongest insolation. But here only shallow roots are directly influenced, and the danger of drought is more serious than that of heat.
Muromtsew (1962) pointed out the fact that plant species have a different amplitude of temperature for root growth. Citrus, for instance, belongs to a group with a small amplitude (7°C), whereas strawberries have a wider one (16°C). This seems to be related to the normal climatic temperature amplitude of the indigenous region. Therefore it is to be expected that trees from the tropics and subtropics should have a narrow amplitude, and trees from temperate and cold regions a progressively wider amplitude. At present, exact and comparable values are lacking. But it seems possible that site tolerance of a species can be limited by the soil temperature, which is especially important in the cultivation of exotic trees. Insufficient root activity as a consequence of low soil temperatures could be a reason for the natural tree line in the alpine and northern regions, because plants suffer from desiccation due to high transpiration and limited water uptake (Michaelis, 1934).
It should be mentioned that temperatures induces morphogenetic changes in roots. Isolated roots of Robinia show an inhibition of lateral root formation at 19°C and an inhibition of length growth of the main root at 33°C, which favors side-root development (Seeliger, 1959). Similar results were obtained by Slankis (1949) with Pinus silvestris. Nightingale (1935) observed that roots of peaches were glistening white with large-diameter, succulent, and fragile tips when grown at 24°C. Roots of Sequoia seedlings appeared to be the healthiest, but not the longest, at 18°C. They were short and thick at 8°C, and thin with fewer and shorter white root tips at 28°C (Hellmers, 1963).
2. ROOT GROWTH-SOIL MOISTURE AND SOIL AERATION Besides soil temperature, soil moisture influences root growth considerably. Useful growth measurements are complicated by the fact that the water uptake by an individual root does not determine its growth rate. A sufficient water uptake by a part of the root system can provide the necessary water for the whole system. Therefore some roots of a system can grow through dry zones when an internal water supply is guaranteed by water uptake of other roots (Shautz, 1927; Kausch, 1959). Because of the unequal distribution of moisture in the soil, this fact is of considerable ecological significance. Of course, mineral salt absorption is inhibited in dry soil zones, as was demonstrated by Hunter and Kelley (1946) with P32.
Water deficiency and water surplus, both have a large influence on the formation of the root system and its activity. In general, root systems are plastic enough to adapt to slowly changing soil water conditions (Rubner, 1960), but rapid changes of the ground-water level can lead to serious root damage because it takes some time before an older root system can adapt to new conditions (Heikurainen, 1964). In soils with high ground-water level we find a "surface," "disk," or "pancake" root system, not only in trees with a natural tendency to form these (Picea abies) but in most other species also.
a. Water Deficiency. Experimental investigations on the influence of soil moisture on root growth have been very infrequent. Therefore nothing is known about specific limit values. Reed (1939) found no significant connection between soil moisture and root growth and stated only generally that growth decreased at low moisture contents. According to Kaufman (1945), daily length growth of Pinus banksiana roots decreased from 3.2 mm (July) to 1.2 mm (August) as a consequence of the diminishing of the soil water from 11% to 2% of capacity. According to Ladefoged (1939), root growth stops in most species when soil moisture is reduced to 12 or 14% on an oven-dry soil basis (or 4 to 6% on an air-dry soil basis). An increase of moisture content above 40% induces almost no additional growth increment (Fig. 11).
In dry soils, roots have a tendency to grow toward more humid zones, so that root enrichments are observed in such zones. In dry soils roots are found at greater depths than in moist soils (Jocum, 1937; Polanskaja, 1962). In silvicultural plow-furrow plantings, fine roots of young pines are restricted to the furrows, presumably because of their higher water content (Buchholz and Neumann, 1964). On the other hand, roots avoid areas of excessively wet soil (Howard, 1925). Roots in swamps lie near the surface, and root growth in depth depends on changes in the groundwater level (Busarova, 1961).
In dry soils the root system has a higher portion of the total plant weight. With increasing soil moisture content the root portion decreases in favor of the above-ground organs (Tolsky, 1904; Aaltonen, 1920; Huber, 1924; Rogers and Vyvyan, 1928; Volk, 1934). In drier soils not only is a larger soil volume made accessible for water absorption by an extensive root system, but the chances of roots encountering scattered local areas of higher water content are also increased. The root system can begin growth anew during drought periods if parts of the system are wetted, thereby increasing the internal water supply to all the roots {Bormann, 1957; Kausch, 1959). Water deficiency leads to an inhibition of root growth before cessation of shoot growth or any visible injury becomes evident (Rogers, 1935; L. M. Turner, 1936; Ladefoged, 1939; Leyton and Rousseau, 1958; Kokhno, 1959). Figure 12 shows this behavior for Larix leptolepis in Germany during the drought of 1964.
Root suberization is accelerated in dry soil and the effective absorbing surface is thereby diminished, so that root systems do not regain their full capacity for water uptake rewetting until some regeneration of growing tips has occurred (Kramer, 1950). When the soil becomes very dry, parts of the root system may die. This is common in surface soil layers (Buchholz and Neumann, 1964). Therefore water and mineral salt uptake remains diminished even for a long time after conditions return to normal following a severe drought. This, in turn, retards root regeneration by inhibition of photosynthesis and brings long-lasting growth depressions in older stands.
b. Water Surplus. Some sites suffer permanently (swamps and peat bogs) or intermittently (Hood plains) from an excess of water in the soil. Water excess-especially in connection with slow water movement- implies a low oxygen availability in the soil. As the results of artificial water cultures demonstrate, tree roots are in general not sensitive to water saturation of the medium, provided that sufficient aeration is maintained. However, ordinarily water saturation of the soil results in a deficiency of oxygen and an enrichment of carbon dioxide, which shifts the redox potential. Furthermore, wet soils are apt to be cold. This lowers the mineral salt and water uptake (Kramer and Kozlowski, 1960); therefore, Hooding or raising of the ground-water level inhibits growth or induces dieback in susceptible tree species as an indirect consequence of an insufficient root activity. Symptoms often first appear during a subsequent drought period, when water supply to the shoot by the partly dead or injured root system is inadequate. Dieback of tree stands in large areas may set in when, because of oxygen deficiency, reduced compounds are formed in the soil which poison the roots (H2S from peat, for example) (Trenel, 1932). On peat bogs, as well as on frozen soils, severe symptoms of mineral salt deficiency are common, although often mineral salt content of the soil is not particularly low.
Roots in swamps are relatively long and poorly branched. After draining of the soil, length growth diminishes, branching increases, and sinker roots are formed which reach greater depths (Heikurainen, 1964). In soils of low moisture content fine roots are usually densely branched.
c. Soil Aeration. Excess water in the soil causes reduced gas exchange between soil and atmosphere so that, by the respiration of soil microorganisms and roots, oxygen content decreases and CO2 and other metabolic products increase. Whether decrease of oxygen or increase of CO2 is more important for root growth is judged differently by various authors. Considerable differences between the tree species probably exist. Boynton ( 1940) mentioned that at 5% CO2 new formation of roots of apple trees is so strongly disturbed that shoot growth is depressed.
Negative effects on growth of agricultural plants by only 1 to 2% CO2 in the soil atmosphere were described by Lundegardh (1957). On the other hand, astonishingly high CO2 concentrations can be tolerated by tree roots if an adequate oxygen supply is provided. This may be as high as 45% CO2 in Salix according to Cannon and Free (1925), or 15 to 60% in cotton according to Leonard and Pinckard (1946). Because of such tolerance, Voigt (1962) considers oxygen the more important factor. According to Leyton and Rousseau (1958) the oxygen requirement is different from species to species (Fig. 13). In Picea nigra even differences between provenances could be demonstrated.
Root growth in Pinus and Picea is noticeably inhibited at values around 10% O2. Cannon and Free (1925) regarded 8 to 10% O2 as minimal for good root growth. In Malus, root growth is already detectably reduced at 15% O2; at 3 to 5% poor growth still can be observed; and at 0.1 to 3% the minimum is reached (Boynton, 1940; Boynton and Reuther, 1938). Only a few species can grow when available oxygen falls below 2% of the soil atmosphere (Fig 13). In the absence of oxygen, roots die after some time. Growth periodicity is of importance in so far as the oxygen requirement during dormant or inactive periods is lower and tree roots are less sensitive. Salix, Alnus, Betula, and some other genera probably are able to provide their roots with some oxygen through an intercellular space system, enabling these species better to tolerate sites poor in soil oxygen (Huikari, 1954; Leyton and Rousseau, 1958). This principle is extremely effective in water and swamp plants which have a large intercellular space system (aerenchyma). In the environs of roots with an internal oxygen supply, heavy metal sulfides in the soil are reoxidized.
Trees are limited in their ability to colonize frequently Hooded or swampy soils mainly by their relative ability to maintain root growth and adequate root metabolism under conditions of poor oxygen supply and lower temperatures (Barner, 1954, 1965). Some trees have developed organs especially adapted for gas exchange in roots (Sonneratia, Bruguiera, and Taxodium species). According to Coster (1933) the resistance of tropical trees to oxygen deficiency in the soil is highly variable. Roots of deep-rooting species are said to have lower oxygen requirements, and oxygen requirements of roots may be an important factor in competition in the tropics (Eidmann, 1935).
3. ROOT GROWTH AND MINERAL NUTRITION
Mineral nutrition also influences root growth and root morphology. In spite of numerous accounts in the literature, it is seldom clear whether the effects described are unspecific and resulting from general promotion of plant growth (for example, as a consequence of fertilization), or whether there are specific influences of mineral nutrition on root growth {Mengel, 1965). The latter have been indicated in some instances, but in general it must be observed that mineral nutrition is closely related to other growth factors such as water supply and shoot activity, and effects of mineral nutrition per se are difficult to ascertain.
It is well known that root development in poor soils is comparatively stronger than in rich ones (Schwarz, 1892; Busgen, 1901; Rogers and Vyvyan, 1928). The root-to-shoot ratio may be nearly 1 : 1 in poor soils, whereas it decreases in better soils to about 1 : 2. Zottl (1964), therefore, with only moderate simplification, states that in vigorously growing stands increased increments of stem wood after fertilization are obtained with unstimulated or only slightly increased root systems. On the other hand, in nutritionally poor stands, additional growth of above-ground parts requires a strong enlargement of the root systems.
A striking fact is the concentration of fine roots in nutrient-rich zones of the soil. Thus stimulated root growth in soil layers rich in humus is often recorded in the field (Moller, 1903; Albert, 1928; Wagenknecht, 1941; Grunert, 1955; Hausdorfer, 1959). In part this may be a nitrogen effect, as it is particularly evident in soils poor in nitrogen (Ehwald et al., 1963). But evidently other nutrients also act in a similar way, because the same phenomenon can be observed in layers of coarse sand with high contents of silicates and in strata of clay or heavy minerals (Fig. 14). A direct influence of soil moisture, although sometimes strengthening the effect, may be excluded. This behavior is caused by the general reaction norm of the root system, which in a state of nutrient deficiency primarily forms poorly branched long roots ("seeking or "pioneer" roots).
In soil layers rich in nutrients, the growth of the main root is lessened at the expense of a strengthened development of side roots, resulting in dense root development in these layers (Fig. 15). This antagonistic behavior between length growth and side-root development is a characteristic feature (Kausch, 1959) which results in ecologically useful modifications of root systems under the influence of certain site factors. Furthermore, it explains the observations that the rooting quotient {total root length/number of root tips) of orchard trees is very high in sandy soils and is reduced after fertilization (Otto, 1964). Various clones, however, may react somewhat differently in this respect. Lundegardh (1957) suggested that the favored length growth induced by nitrogen deficiency be called "nitrogenium-deficiency-etiolement."
In addition to the above, it must be noted that the mineral nutrient status (and especially nitrogen supply) affects the root-to-shoot ratio. Therefore, in the field, differences occur in intensity of rooting in various humus forms with different nitrogen contents (Table IV). In raw humus poor in nitrogen, Fagus silvatica forms about twice as much length and weight of roots per square centimeter of leaf area as in mull (Meyer, 1963).
Humic acids and quinoid compounds have been reported to elicit specific plant growth effects. Stimulation of root growth has also been reported (Flaig, 1958; Giulimondi, 1961). Little is known about the possible effects of other biogenic compounds formed in the soil on root growth and development.
Specific effects of different nitrogen compounds are noteworthy. Leyton (1952), Prjanischnikow (1952), and Smith (1957) found that the root systems of plants supplied only with nitrate are stronger than of those plants fertilized with ammoniacal nitrogen. Evers (1964) also considers nitrate to be the best form of nitrogen for growth of poplar. Alnus glutinosa fertilized with nitrate develops very dense, fine, and abundantly branched root systems, whereas ammoniacal fertilization causes formation of long and sparingly branched roots. Under field conditions there is probably a relation between soil aeration and the form of soil nitrogen.
An extensive and rather contradictory literature exists concerning the effects of other elements on root growth and the formation of root systems. Because of ion antagonism and multifactorial effects which may lead to variable results in different cases, valid generalizations for woody plants are not yet possible. (For further literature see Kramer, 1956.) Fertilization of forests generally stimulates superficial rooting because vertical translocation (especially of phosphorus) is rather slow. Furthermore, the humus content and the recycling of nutrients by soil organisms favor the enrichment of nutritive elements in the upper soil horizons. Buchholz and Neumann (1964) found that in a 56-year-old pine stand the superficial rooting was doubled two years after nitrogen fertilization, while at the same time the deep rooting decreased (Fig. 16).
4. ROOT GROWTH AND LIGHT INFLUENCE The formation of the root system is dependent-as is plant growth as a whole-on the photosynthetic efficiency of the tree, which means that root growth is in competition with shoot growth for carbohydrates. In higher plants a general and ecologically useful reaction norm in the distribution of carbohydrates has evolved, especially when shading makes
carbohydrates limiting. Increasing shade decreases growth as a whole but leads to a relative stimulation of shoot growth at the expense of root development (Mitchell, 1936; Gast, 1937; Kozlowski, 1949; Lyr et al., 1963). In short, shading primarily influences root growth, and the root-to-shoot ratios are thereby altered.
The intensity of the effect, however, varies according to the shade tolerance of the species involved. From this it is clear that at reduced light intensities the ability to wage root competition decreases in shade- intolerant species. This effect may be reinforced by other factors that
induce a relative reduction of root growth-for example, nitrogen fertilization (Lyr et al., 1967). Shaded plants are therefore apt to be more susceptible to drought than others in full light (Kramer and Decker, 1944; Barney, 1951). At a light reduction to 40% of daylight, which is by no means an extreme shading degree under field conditions, both rooting depth and rooting density of Robinia are strongly reduced Whereas plants in full daylight showed a total length of fine roots of 266 m, the corresponding value in the shaded ones was only 39 m (Hoffmann, 1965). In Lupinus and Alnus, shading greatly reduces nitrogen fixation (Hoffmann, 1960; Lyr et al., 1963). Some other tree species show remarkable reduction in formation of mycorrhizae when shaded (Bjorkman, 1942).
W. Turner (1922) and Shirley (1929) earlier pointed out that the strongest root systems are developed in full daylight. The former believed that surplus quantities of assimilates not required for shoot growth are used for root growth. This may be only partly correct, however. If a
sufficient nitrogen supply is provided, shoots compete with roots whenever environmental conditions allow growth. Whether trees of the Populus type, having long duration of shoot growth, and of the Quercus type, which have a short growth period (Lyr and Hoffmann, 1965), behave differently in root/shoot competition has not yet been determined.
Reduction of photosynthetic activity, although caused by shading of the leaves, results in decreasing root growth, but not immediately. The reaction of the roots takes place after 12 to 24 hours in Acer saccharinum (Richardson, 1953a,b). This may be the reason why root growth is more active at night than during the day (Section 1I,B,4). In young Acer seedlings this root reaction is more rapid, whereas in Quercus the reaction of root growth as a consequence of shading takes place more slowly, presumably because of higher food reserves in this genus (Richardson, 1956). The negative phototropism of roots is well known. But various species are likely to exhibit different sensitivity. Isolated roots of some tree species show decreased growth in the light (Seeliger, 1959).
5. ROOT/SHOOT INTERRELATION
Root growth and shoot growth are closely interrelated. Physiological regulation mechanisms of an unknown nature provide for a balanced root-to-shoot ratio adapted to the ecological conditions. Probably hormonal mechanisms which determine correlative food distribution are of great importance. There exist some generally applicable reactions norms, which vary quantitatively from species to species. The root as a heterotrophic organ is dependent on the shoot for a supply or carbohydrate and some vitamins. This can be demonstrated with cultures of isolated roots (Slankis, 1949; Seeliger, 1956, 1959; Ulrich, 1962). The level of food and auxin supply from the shoot to the root depends on the conditions affecting photosynthesis, on leaf age and leaf area, and on the utilization of photosynthate within the shoot. When the later is low-for example, in rooted leaves-a large root system is built up very soon. The reaction of root growth to changes in rate of photosynthesis and respiration is, therefore, understandable (Richardson, 1953a,b; see also above).
Although roots are not dependent on auxin supply from shoots, hormone production of shoots seems to influence root development in a specific manner. According to Richardson (1958) defoliation inhibits length growth of roots but not root-sucker formation. Defoliation of the leading shoot bud inhibits formation of new roots, whereas roots already present go on growing. f3-lndoleacetic acid can counteract the effect of decapitation. Probably still other hormones participate in the regulation of root growth (Romberger, 1963).
Environmental factors and cultural practices can shift the root -to - shoot ratio by incompletely known mechanisms. Soil dryness, mineral salt deficiency (especially of nitrogen), and higher soil temperatures cause an increase in ratio, whereas shading, nitrogen fertilization, higher air temperatures, and sufficient soil moisture induce a decrease in the root-to-shoot ratio. Pruning, crown cutting, and defoliation act like shading and inhibit the development of the root systems (Chandler, 1923; Heinicke, 1936; Sawage and Cowart, 1942; Wood, 1939; Hoffmann, 1966c).
Mowing and grazing of herbaceous plants decreases their root formation and the ability of their roots to compete (Weaver and Darland, 1949; Kohnlein and Vetter, 1953). Development of the root system may also be reduced by heavy fruiting. Chandler (1923) found a 50% decrease in the root system of Prunus after a heavy crop, and, according to Nutman (1933), Coffea arabica can be injured or killed by high harvests because of the insufficient supply of carbohydrates to the root system.
Some data indicate that typical species-to-species differences exist in the root-to-shoot ratio. This has been investigated mostly in young trees.
According to Lobanow (1960), the ratio of root surface to leaf surface is less than 1 in strongly mycotrophic trees (such as Pinus, Picea, Larix), whereas nonmycotrophic species reach much higher values. A value of 139 has been reported for rye (Dittmer, 1937). Mycorrhizae probably influence the root-to-shoot ratio by providing additional absorbing organs.
In older trees it is assumed that about 20% of the total weight is roots (Rogers and Vyvyan, 1934; Ehwald, 1957; Assmann, 1961). According to Bray (1963) and Whittaker et al. (1963), root-to-shoot ratios decrease with increasing age of trees. The above-ground and subterranean development of a tree are always in close correlation. The more exactly the real assimilation efficiency and the root quantity are ascertained, the better can this correlation be determined. The crown radius, for example, gives a lower correlation than the crown mantle area (Melzer, 1962a).
The root-to-shoot ratio changes during the growing season because of the somewhat independent development of roots and shoots. This is evident in Fig. 8. Trees having a Quercus type of growth flush show this most distinctly. A typical course for Picea glauca (Mullin, 1963) is shown in Fig. 20.
Finally we want to mention some peculiarities. According to Lemke ( 1955), clear correlations exist in Quercus borealis var. maxima between crown size and trunk diameter on the one hand and the mean root depth and range of horizontal roots on the other. Large crowned trees have a denser root systems than others. In contrast to other trees, the horizontal roots of Pseudotsuga taxifolia reportedly do not reach beyond the crown projection area (Wagenknecht, 1958), which is important from the silvicultural viewpoint. In this same species a one-sided crown development is associated with a one-sided formation of the root system.
The management of stands and trees for good crown formation is a possible way to assure the production of large, deep-reaching root systems which give a sure protection against windthrow (Wagenknecht and Belitz, 1959). Whereas in general dominant trees develop a deeper root system than suppressed ones, in Fagus silvatica root depth is quite independent of the socia-ecological position of the tree (Wagenknecht, 1960).
It is interesting that the formation of a new leading shoot from a flat-topped crown in Pinus silvestris is correlated with the regeneration of a new taproot (Albert, 1907; Wagenknecht, 1960). According to Romer and Hilkenbaumer (1936, 1937) scions also have a specific influence on growth rate and branching of roots of orchard trees. Similar observations were made by Kemmer (1964).
D. Practical Considerations
In efforts to increase output of forest products by application of a highly developed forest science, there is a need for exact knowledge of the causal connections between environmental effective factors and tree response. In silvicultural decisions, knowledge of root growth and the factors influencing it should play a more important role than it has in the past.
It is possible, by numerous practical steps, to influence root growth and development of the root system in order to increase tree growth and to protect forests against harmful biotic and abiotic agents. There are data indicating that it is likewise possible by breeding to select clones or populations of trees which have the desired qualities with respect to root formation and root activity.
Some management measures based on knowledge of root behavior are already being applied in the forest. It is well known, for instance, that crown-tending diminishes the danger of wind throw on wet or shallow soils where trees have a tendency to form superficial root systems. There is a fund of practical experience concerning the suitability of various tree species for heavy, difficulty penetrable soils, such as pseudogleys. On such soils, as well as on periodically flooded sites, a high root activity and a low demand for exogenous oxygen are decisive factors in determining site tolerance of a tree species.
Tree species in mixtures that have proved to be highly productive are often complementary to one another in the utilization of the site by the root system as well as in utilization of light. Therefore mixed stands often are more productive than pure stands (Erteld, 1953). New mixtures and mixture ratios can be developed theoretically from the knowledge of the general physiology of the trees and of the behavior of their root systems. This is very important in the tropics, where root competition is often a dominating factor.
Soil temperature has often been left out of consideration as a site factor, although it has been demonstrated in several instances that an increase of soil temperature brings an increase of yield. The failure of some trees on cold soils is caused by an insufficient root metabolism which causes deleterious effects because of an inadequate supply of water, mineral salts, and essential organic compounds to the shoot. The periodicity of root and shoot growth as well as the varying ability to regenerate roots during the vegetation period should be considered in the choice of planting times and times of tending and fertilizing young plantations.
Preparation of the soil should be done in such a manner that sufficient aeration is guaranteed and a deep-reaching, drought-resistant root system is formed. This may be achieved on some soils by a suitable layering of the humus by plowing.
Fertilization should be done in such a way that a harmonious root-to- shoot ratio is maintained. This is important in nurseries to minimize losses arising from climatic anomalies. Development of a shallow root system as a consequence of fertilization should be counterbalanced by other measures specifically intended to promote deep rooting.
Light conditions in stands can be influenced so that strong root systems are developed which have a high ability for competition and for utilization of deeply situated water and mineral salt sources. Here differences between tree species should be observed and considered in selecting species combinations in mixed stands.
Many failures of cultural techniques as well as growth stagnation of trees and stands have their cause in poor root growth or in damages to the root system. Their early recognition and amelioration are important factors for increasing production. The rate of regeneration of the root system, which can be favored by several practical measures, often determines the duration and severity of growth interruptions and losses of increment.
Further examples of possibilities of increasing wood production by practical application of knowledge about root growth and root behavior could be given. At present, unfortunately, detailed data on quantitative differences in the behavior of tree species and their provenances are still lacking. Available observations are too sporadic to permit valid conclusions to be drawn from them. Therefore, reviewing the literature and summarizing the facts and opinions found therein is not easy. But in our endeavors we should not shrink from attempting to analyze the complicated physiological processes of interaction with the environment and from striving to build up a theoretical base for a high silvicultural production.
REFERENCES
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50. The Rhizosphere 

One of the first articles that really made me aware of this amazing world beneath our feet. It is written without knowledge of glomalin but comes right up to the edge of it without seeing it. Also critical is the understanding of forests as systems and trees as pumps. By now commercial mushrooms are an insignificant itemas the fungal processes of their lives come into the spotlight. This article was posted on the John A. Kelsik and Son web site.

Rhizosphere Wars
The rhizosphere is the absorbing root-soil interface. It is the zone, about one millimeter in width, surrounding the epidermis of living root hairs and the boundary cells of mycorrhizae as well as hyphae growing out from some mycorrhizae.
The rhizoplane is the boundary where soil elements in water are absorbed into the tree. Under an electron microscope, the rhizoplane appears as a jelly where microorganisms and tree cells mix, making it impossible to tell which side is tree and which is soil.
A constantly changing mix of organisms inhabit the rhizosphere and surrounding soil. Bacteria, actinomycetes, fungi, protozoa, slime molds, algae, nematodes, enchytraeid worms, earthworms, millipedes, centipedes, insects, mites, snails, small animals and soil viruses compete constantly for water, food, and space.
The rhizosphere is a battleground and the wars are continuous. Amoebae are eating bacteria. Some bacteria are poisoning other bacteria. Fungi are killing other fungi. Nematodes are spearing roots. Fungi are trapping nematodes. Earthworms are eating anything they can find. Sometimes the victors benefit the tree and sometimes they do not.
Every tree treatment affects the rhizosphere in some way. The more you know about the rhizosphere, the better the chances are that your treatments will lead to benefits rather than harm.
Declines and the Starving Rhizosphere
Go anywhere in the world and you will learn that some local trees have a "new" decline problem. Declines usually mean the trees are sick because there is a problem in the rhizosphere.
Trees die, as all organisms do, in three basic ways: depletion, dysfunction and disruption. Disruption means wounding, severe mechanical impacts and fracturing. Dysfunction means some parts and processes of the living system have developed problems that retard or prevent their functioning and growth. Depletion means that the basic substances for life begin to decrease to the point where injury and death are certain. One of the ways depletion injures organisms is by starvation.
Soils and wood share a common problem: They are thought of as dead substances. This has come about because wood-products research gained an early lead over research on wood in living trees. With soils, many texts still define soils as "loose material of weathered rock and other minerals, and also partly decayed organic matter that covers large parts of the land surface on Earth."
Sapwood in living trees has many more living cells than dead cells. In upper layers where most absorbing roots of plants grow, soils have more soil organisms than grains of weathered rock. In great disrespect, most people still refer to soil as dirt! When researchers first discovered the great value of soil microorganisms for human antibiotics and profit, the living nature of the soil began to emerge.
A more correct definition of soil should be that it is a substance made up of sands, silts, clays, decaying organic matter, air, water and an enormous number of living organisms. Survival of all living systems depends greatly on synergy and efficiency to optimize the functioning of all processes and to keep waste as low as possible. When synergy and efficiency begin to wane, declines follow.
Trees are dependent on the light energy from the sun for their energy, water and 14 elements from the soil for their building blocks of life. Some trees decline when incorrect treatments or abiotic injuries lead to starvation of organisms in the rhizosphere. When there are troubles in the rhizosphere, there will be troubles with the tree.
Energy & Root Exudates
Microorganisms compete in the rhizosphere, an area rich in exudates from the tree. The exudates contain carbohydrates, organic acids, vitamins and many other substances essential for life. From 5 percent to 40 percent of the total dry matter production of organic carbon from photosynthesis may be released as exudates! When trees begin to decline, the amount of organic carbon released as exudates increases. Mineral deficiencies, low amounts of soil air and severe wounding are major causes for the increase. Another way to say this is that an increase in exudates would be caused by over-pruning, construction injury, planting too deeply, over-watering, compaction and planting trees in soils that have a pH too high or too low for their optimal growth.
You would think that a tree in decline would decrease not increase exudates. A possible explanation might come from the self-thinning rule of ecology, which states that when energy input into a site equals output, there will be no further growth unless some trees die. As many suppressed trees die, a much fewer number continue to grow bigger. Simple. Or, on the basis of the mass-energy ratio law, as some trees on a site get bigger, many smaller suppressed trees will die. As the suppressed trees decline, they contribute a higher percentage of their soluble carbohydrates to the rhizosphere.
The increase in exudates from a declining tree with a defense system weakened by low energy reserves would give root pathogens an advantage over other soil organisms. When the tree dies, its dead wood adds a great amount of carbon to the soil, thus benefitting all soil organisms. If this scenario is correct, then the codes for the increase of exudates as trees decline would have been set in the genes of the forest trees. Then, even after trees are taken out of their groups in forests and planted as individuals, the genetic codes for increasing exudates as the tree declines for reasons other than crowding would still be in effect.
A tree does not "know" why it is dying. In a crowded, young, growing forest, the self-thinning rule of ecology does benefit tree survivors and all soil organisms. But, when one or two trees in a yard, city or park start to decline, their early death may benefit only the root pathogens. And even worse, since the tree will be cut and removed from the site, there would be no benefits from added carbon to the soil.
A Closer Look at Roots
Woody tree roots are organs that support the tree mechanically, store energy reserves, transport water and the substances dissolved in it and synthesis substances such as growth regulators, amino acids and vitamins that are essential for growth.
Trees have different types of root systems. For example, mangroves along coastlines have stilt roots. Many trees growing in tropical areas have aerial roots that become prop roots when they grow into the soil. Other trees have strangling roots that eventually kill the host tree that first supported their growth. Trees in sandy soils can have roots that grow downward over 90 feet. Palms have roots that are adventitious and grow from meristematic regions in their base. Many tree species have deep roots when they are young and more shallow roots later. It would be nearly impossible for the strongest person to pull out young saplings of beech, oak or hickory from forest soil.
Woody roots have cells with walls of cellulose, hemicellulose and lignin. Lignin is that natural "cementing" substance that gives wood its unique characteristic for strength. Woody roots also have an outer bark or periderm made up of three layers: the phellogen, phelloderm and phellem. The phellogen is the bark cambium. The phelloderm is a thin layer of cells on the inner side of the phellogen. The phellem is the outer corky layer. Phellem cells are impregnated with a substance called suberin, which is a fatty substance that prevents water absorption.
Some characteristics of woody roots are:
* They do not absorb water.
* They have no pith.
* Their conducting elements are usually wider than those in the trunk.
* They have a greater proportion of parenchyma cells than is usual for trunks. The living parenchyma store energy reserves, usually as starch.
A soft cortex without chlorophyll may be in the bark. In some tree species that thrive in wet soils or have deep roots, the cortex may have many open spaces that act as channels for air to reach the living cells in the roots. It is important to remember that the parenchyma in the woody roots store energy reserves, and root defense is dependent on energy reserves. When reserves are low, defense is low. When defense is low, weak or opportunistic pathogens attack. It is nature's way.
Non-Woody Roots
Non-woody tree roots are organs that absorb water and elements dissolved in it. The two basic types of non-woody roots are:
1. Root hairs on non-woody roots are extensions of single epidermal cells. Common on seedlings, root hairs grow to maturity in a few days. They function for a few weeks and then begin to die.
On mature trees, they are usually not abundant. When they do form, they do so when soil conditions are optimum for absorption of water and elements. I have found root hairs growing in non-frozen soils beneath frozen soils in winter.
2. Mycorrhizae are the other type of non-woody roots. Mycorrhizae are organs made up of tree and fungus tissues that facilitate the absorption of phosphorus-containing ions and others essential for growth.
The fungi that infected developing non-woody roots to form mycorrhizae were very "biologically smart." Rather than competing with other microorganisms in the rhizosphere for exudates from the tree, the mycorrhizal-forming fungi went right to the source inside the tree. And, even more to their advantage, many of the mycorrhizal fungi grew thread-like strands of hyphae-long, vegetative tubes of fungi-out from the mycorrhizae. This inside and outside presence gave the fungi a distinct advantage over other microorganisms in the rhizosphere.
The tree gains efficiency with mycorrhizae in several ways.
1. With their extended hyphae, mycorrhizae not only greatly extend the absorbing potential into the soil, but the hyphae may connect with other hyphae on other trees. In this way, the mycorrhizae serve to connect trees of the same or a different species. This leads to the conjecture that the natural connections that developed over long periods in the natural forest may have some survival value. That is why forest types are often named for the groups of species commonly found growing together. For example, we speak of the birch-beech-maple forest, or the pine-oak forest. From a practical standpoint, when trees are planted in cities and parks, there may be great survival advantages by planting groups of trees made up of the species that are normally found together in natural stands.
2. The mycorrhizae have been shown to provide some resistance against root pathogens. It may be that the pathogens would have difficulties in building their populations in the rhizosphere dominated by the mycorrhizal fungi.
Perhaps the most important feature of the mycorrhizal fungi is that their boundary material is mostly chitin. Chitin is slightly different from cellulose by the replacement of some cellulose atoms by a chain of atoms that contain a nitrogen atom. This slight change in some way makes chitin a material better suited for absorption of elements. Remember that the fungus hyphae gain all their essentials for life by absorption through their boundary substance.
There are other advantages, to the chitin and the tube-like hyphae that ramify the soil in the rhizosphere and beyond. When the hyphae die, they add a nitrogen source for other organisms. Also, when the hyphae are digested, they leave tunnels in the soil that are about eight to 10 microns in diameter. For the bacteria, these small tunnels may mean the difference between life and death. The bacteria quickly colonize the tunnels. The survival advantage here is that the major threats to their survival are protozoa that are usually much larger than 10 microns. So the hungry amoebae are not able to get at the bacteria inside the eight-micron tunnels.
A common treatment for compaction is to fracture the soil and add water,. The fracturing allows air to penetrate the soil, but does not provide any eight-micron tunnels for the bacteria. The only way to bring back the tunnels is to bring back the fungi in well-composted wood and leaf mulch, as nature does, or by inoculating the mulch with mycorrhizal fungi.
Who Was First? I do not know if the fungi were the first to grow into the root to get first chance at exudates or whether it was the bacteria. Regardless, bacteria and their close relatives, the actinomycetes, also infect non-woody roots to form organs that serve for the fixation of atmospheric nitrogen. Fixation means that the nitrogen that makes up almost 80 percent of our air is converted to a soluble ionic form by the action of the bacteria and actinomycetes within the nodules on the roots. (Some free-living soil bacteria can also fix nitrogen.) An enzyme called nitrogenase is the catalyst for the reaction that will take place only under very exacting conditions. There must be soluble molybdenum and iron and no free oxygen available. These conditions are present within the nodules. Here again, the microorganisms benefit the tree by providing a source of soluble nitrogen, and, in turn, the bacteria and actinomycetes get first chance at exudates. Even more importantly, the nodules protect them from foraging protozoa.
Infections that result in benefits to both parties are called mutualistic. When the benefits are greater than the sum of the parts, the association is called synergistic.
Species of legumes commonly have bacterial nitrogen-fixing nodules and mycorrhizae. The mycorrhizae facilitate absorption of elements, and the nodules provide a nitrogen source. Many species of trees have actinorhizae, which are the nodules formed by the root infections by actinomycetes. Species of Alnus have very large nodules. The actinorhizae are common on tropical and subtropical trees, and especially on trees that have adapted to soils low in available elements essential for life.
On some subtropical and tropical trees, such as the macadamia, multi-branched clusters of non-woody roots called proteoid roots form. The proteoid roots alter the rhizosphere by acidification processes that facilitate the absorption of phosphorus-containing ions. When I examined the roots of dying macadamia nut trees in an orchard in Hawaii, I could not find proteoid roots, yet only a few days earlier I had found them on macadamia nut trees growing in the wild. I learned later that the orchard where trees were dying was heavily fertilized on a regular, basis with phosphorus.
Another type of nodule forms on species of cycads. These nodules harbor blue green algae, or cyanobacteria, that have the ability to fix atmospheric nitrogen.
My point is that many different synergistic associations have developed in, on and about non-woody roots that provide elements, not an energy source. These associations are of extreme benefit to all connected members. At the same time, the conditions that provide for the associations are very delicate and exacting. It does not take much to disrupt them.
It Does Not Take Much to Disrupt Them
This statement deserves repeating and repeating. The delicate "threads" that hold these powerful associations together need to be recognized and respected. Trees in cities grow only so long as these "threads" remain connected.
Trees grow as large oscillating pumps, with the top trapping energy and pumping it downward. The bottom absorbs water and elements and pumps them upward. The pumps have developed over time to work on the basis of many synergistic associations that maximize benefits for all connected members and to minimize waste.
Many of life's essentials for the bottom associates come from the top of the tree. And, the top works only because the bottom works. Energy is required to move things, and elements and water are required to build things.
Tree Treatments and the Rhizosphere
When trees are over-pruned, the top will be injured first. When it is injured, it will not serve the energy requirements of the bottom. Soon root diseases start and are blamed for the decline or death of the tree. Where over-pruning is common. so are root diseases.
Compacted soil blocks air and water to the bottom and crushes all the microcavities where the microorganisms live. In nature, decomposing wood and leaves keep conditions optimal for the rhizosphere inhabitants.
Over-watering stalls the respiration processes in the roots. When respiration stops, carbonic acid is not formed. When carbonic acid is not formed, ions necessary for the absorption process do not form. When absorption is down, the tree system is in trouble. Fertilizers can be of great benefit to trees growing in soils low in or lacking elements essential for growth.
Elements or molecules made up of a few to many different atoms enter the roots as ions. An ion is a charged atom or molecule. Ions with a positive charge are cations, and those with a negative charge are anions. Each particle or granule of fertilizer is a salt made up of a lattice of anions and cations, just as ordinary table salt is made up of a grand lattice of connected sodium cations and chloride anions. When salt as sodium chloride dry granules is poured into water, the sodium and chloride ions separate. When they separate, they carry electrical charges and are called the sodium ion and the chloride ion. When a cation enters a root, another cation must exit. This is very important, as we will see. When nitrogen enters a root as nitrate anion, an anion of bicarbonate ion from carbonic acid exits. The bicarbonate ion is probably the second most important compound in nature, next to water, because it drives the absorption process. When a bicarbonate ion exits into the rhizosphere, the pH increases.
When urea is used in fertilizers as the nitrogen source, the pH in the rhizosphere could increase to 2 or more pH units. The chemistry behind this is complex, but here I present only the conclusion, because a common problem with trees in some high pH soils is chlorosis. There is no easy field method for measuring the pH of the one millimeter wide rhizosphere. The rhizosphere could be pH 8, and the bulk soil would measure pH 6. As pH increases, the availability of elements such as iron and manganese decreases. In soils, it is one thing to have an element present and another to have it in a form available to the plant as an ion. As pH increases, iron and manganese element, form molecules that precipitate in water rather than ionize. If they are not available as ions, they will not be absorbed. And, if they are not absorbed, several of the enzymes essential for chlorophyll formation and photosynthesis will not form.
When the energy flow from the top of the pump is blocked, then the bottom does not get enough energy for growth and defense. The pathogens invade, and the tree declines. This scenario does not mean that every time you use urea, trees will decline from chlorosis. But the use of urea could be a contributing factor where trees with genetic codes for growth on low pH soils are planted in high pH soils. If fertilization is a desired treatment, then a fertilizer that has nitrogen in a positive charged ion, such as an ammonium ion, would help to reduce the rhizosphere pH. When the ammonium ion enters the root, a proton of positive charge will exit. The protons in rhizosphere water will bring about more acidic conditions, so there is a way out.
In summary, fertilizers can be very beneficial for healthy survival of trees planted outside their forest homes. How beneficial will depend greatly on an understanding of many of the points mentioned here and some basic chemistry.
Primary Causes of Diseases
It is often very difficult to have people recognize the importance of small organisms in small places doing big things. Blame for the death of a tree is often placed on big things that can be seen or felt. Most pathogens are opportunistic weaklings waiting for a defense system to decrease. Many small disrupting events often lead to the decrease in a defense system. Then after the tree has been weakened, the final agent comes along and gets the full blame for the cause. A perfect example is the cankers on honey locust. Flush pruning is usually the real cause.
Pumps and Food Trees are oscillating pumps. When the pump begins to wobble, some parts will begin to weaken. When they weaken to the point where some other agent causes a part to break, the pump will stop.
It is very difficult to determine where problems start in an oscillating pump. Symptoms may be in the bottom, but the cause may have been in the top. Or, it could be the other way around.
I go back to two points that may be part of the answer: exudates and the self-thinning rule of ecology. All living things require food and water for growth. Leaves and photosynthesis provide the energy at the top of the pump. The nonwoody roots and the rhizosphere provide the elements and water at the bottom. Photosynthesis will not work without water and elements, and the absorption processes will not work without an energy source.
Trees became trees growing in groups in forests where the self-thinning rule had strong survival value. Not only did exudates provide quick energy for the rhizosphere organisms, but the carbon in the wood of the trees that fell to the ground also provided a long-lasting energy source for a succession of organisms.
Reports from some countries indicate an abundance of soluble nitrogen compounds in runoff water and even in ground water. This is a strong indication that the carbon-nitrogen ratio has been disrupted in the soil. It is well established from studies of the physiology of fungal parasitism that the degree of parasitism is often determined by the carbon-nitrogen ratio. It is probably similar for other organisms.
The organisms in the rhizosphere and surrounding soils have many different ways to weather rocks and to get nitrogen and other elements essential for their growth. What they cannot get in the soil is a sufficient energy source. Yes, some small animals die and provide carbon, and some microorganisms can get energy by chemosynthesis, but the requirements for carbon are much greater than what could be supplied by those sources alone. Carbon must come from the top of the pump. When the energy source from the top begins to decrease, the rhizosphere organisms will begin to starve.
The oscillating pump model soon takes on the form of a circle, because now it could be said that the top did not work efficiently because the bottom had a problem first, and this could be so. My point is that the energy problem does play a key role in declines. If a single tree is already very low in energy reserves, it cannot contribute much to the rhizosphere even if the genetic codes rule that exudates should increase as a tree begins to decline. Soon we will be faced with the chicken or egg problem.
I believe there is a way to decrease the potential starvation problem. In forests, more wood should be left on the ground, and in cities, more composted wood and leaves should be added in correct quantities to the soil about the base of trees. Incorrect treatments of pruning, watering, planting and fertilizing should be corrected, because they often start the pumps to wobble. If these simple adjustments can be made, rhizosphere starvation will decrease and our trees will lead healthier and longer lives.
Author's Note
Much of the information presented here has come from several books that I found very helpful in preparing for this article. I recommend these few books to people who want more information.
1. Foster R.C , A. D. Rovira, and T.W. Cock. 1983. "Ultrastructure of the Root-Soil Interface." The American Phytopathological Society, St. Paul, MN.

2. Kilham Ken. 1994. "Soil Ecology." Cambridge University Press. Cambridge, Great Britain.

3 Wild, Alan. 1994. "Soils and the Environment: An introduction." Cambridge University Press. Cambridge, Great Britain.

Dr. Alex Shigo is a noted authority in the field of modern arboriculture. An author, lecturer and consultant, he is the owner of Shigo, Trees & Associates in Durham, New Hampshire.
Reproduced with permission of Tree Care Industry and Dr. Alex L. Shigo.
The article was published in Volume VII, Number 10 -October 1996 of TCI.
This site is dedicated to the remembrance to Robert Felix who for many years worked very hard for the improvement of the tree care industry: 1934-1996.

Please report web sit problems to John A. Keslick, Jr.
Back to articles.

49. Soil Biological Communities 

A good basic understanding of the soil and its inhabitants is available at BLM's Soil Biological Communities. This is the page about mycorrhiza, links there take you to bacteria, nematodes and so forth. Most plants are VA mycorhizzal, most forest trees ectomycorhizzal, also producers of soil glue. They are the majority of the commercial mushroom industry, and of course, responding to elevated CO2.
Mycorrhizal Fungi
What They Are and A Few Interesting Facts

http://www.blm.gov/nstc/soil/index.html
Mycorrhizal fungi colonize the roots of many plants. Mycorrhizal fungi don’t harm the plant; on the contrary, they develop a "symbiotic" relationship that helps the plant be more efficient at obtaining nutrients and water. In return, the plant provides energy to the fungus in the form of sugars.
Here’s how that symbiotic relationship works. The fungus is actually a network of filaments that grow in and around the plant root cells, forming a mass that extends considerably beyond the plant’s root system. This essentially extends the plant’s reach to water and nutrients, allowing it to utilize more of the soil’s resources.
There are two main categories of mycorrhizae common to western rangelands in the United States.
Vesicular-arbuscular mycorrhizae or VAM. VAM is a type of endomycorrhizae (endo = inside), and is the most widespread of the mycorrhizae. These fungi actually reside inside the cells of the plant root. They’re typically found associated with most grasses, forbs, shrubs, and a few trees such as juniper. They are generalists, have only a few species, and are slow to disperse.
Ectomycorrhizae (ecto = outside) grow around the root and between the root cells, but unlike VAM, the fungus doesn’t actually penetrate the root cells. The fungus also forms a considerable mass in the soil surrounding the plant roots. The fruiting, or reproductive bodies, of these fungi are sometimes visible as something we all recognize— mushrooms! Ectomycorrhizae are commonly associated with forest trees of temperate regions. On rangelands, they are found in riparian areas (the places next to water), open woodlands, and shrub oak communities. They are host-specific, have many species, and can disperse far and quickly.
Not all fungi are mycorrhizal. There are also fungi that help decompose the organic matter in litter and soil. However, they play a lesser role than bacteria in this important process in semi-arid and arid rangeland soils.
Why They Are Important
Some plants are "mycorrhizal-obligate," meaning that they can’t survive to maturity without their fungal associate. Important mycorrhizal-obligate plants in western North America are sagebrush, bitterbrush, and some native bunchgrasses.
Mycorrhizae are particularly important in assisting the host plant with the uptake of phosphorus and nitrogen, two nutrients vital to plant growth.
Mycorrhizae actually increase the surface area associated with the plant root, which allows the plant to reach nutrients and water that might not be available otherwise. Put simply, mycorrhizae extends the plant’s reach, allowing it to get to more of what it needs to survive. That makes the plant stronger, especially during drought periods. Stronger individuals means that the community is more resilient to disturbance. Some mycorrhizae may even protect their host plant against unwanted pathogens.
Learn More!

Allen, M.F. 1991. The ecology of mycorrhizae. Cambridge Univ. Press. New York. 184 pp.

Ingham, Elaine R. 1998. The soil biology primer, soil fungi. USDA, Natural Resources Conservation Service, Soil Quality Institute.

Read, D.J. 1991. Mycorrhiza in ecosystems. Experientia 47: 376-391.

Wicklow-Howard, M. 1994. Mycorrhizal ecology of shrub-steppe habitat in Proceedings-Ecology and Management of Annual Rangelands. USDA, Forest Service. Inter-mountain Research Station. General Technical Report INT-GTR-313. pp. 207-210.

Wicklow-Howard, M. 1998. The role of mycorrhizal fungi in rangelands. pp. 23-25 in Rosentreter, R. and A. DeBolt, editors. The Ellen Trueblood Symposium. Technical Bulletin No. 98-1, Bureau of Land Management, Boise, Idaho.

48. Two Days of Culmination 

This week I spent two days in southern Humboldt wwrapping up two long in the works projects. Thursday BLM, MRC,MMC and Humboldt Redwoods State Park Interpretive Center Operator Dave Stockton walked parts of Gilham Butte for writing management options for the Coopereative Management Plan. Public efforts to "Save Gilham Butte" have abounded three times since 1976; it has finally been spared the harvesting saw.
We walked through oldgrowth Douglas fir forests that needed minimal treatmnet. We walked through younger stands of tanoak with good spacing and a closed canopy, a hardwood stand also needing little or no treatmnet. Adding in remote and difficult access, this area has little cost effective needs.
This mangemnt option is where forestry, glomalin, recreation, wildlife and fire risk converge. First we must fix the machinery of the forest, then it should be allowed to run for a while to get back on track. Large old trees fix lots more CO2 than many little ones and so need to be preserved as ongoing operations. This is an operating forest managed for maximum forest efficiency, not a hands off preservation policy. We salute BLM for developing TSI standards for non-harvestable lands like King Range wilderness areas and Headwaters.
Next day I met with Good Roads Clear Creeks coordiator Dylan Brown and site assessor Joel Munschke on my property on Middle Creek. It burned in 1981 and has been subject of a dozen hikes with MRC and others like Scott Downey and MSG since 1987 and peaking with the sediment inventory done in 2001. The work clearly shows that road drainage diversions cause downslope ground failuires. What we have really been after is help with costs and permits for instream improvements. We needed to stabilize the slopes first. About all you can do is roadwork and revegetation.
In the creek itself, several days of excavator work will move the entire creek away from the failing soil bluffs and into an old channel already shaded by conifer regrowth. There is a lot of rock onsite to secure it, and to use all the available money for machine time, rather than hauling rock in. This will vastly improve around 500 yards of salmonid spawning habitat. GRCC expects to accomplish this landscape sized task in the next month.
Here we cannot rest however, revegetation has now become a fire risk. Thinning and release cuts are needed on large scales. Funneling the growing power of the land to restore system functions and lower risk need active management. Land in the ceanothus need replanting with forest trees. Douglas fir are full of cones this year and we will get another round of direct seeding from trees fifteen to twenty years old.
We salute MRC for this ambitious project, and its cookie cutter repeatability. These treatments, on a one time around basis, coupled with time and natures healing ability, should accelerate reccovery to the point of landscape stability even in heavy precipitation events.
Long-Term Effects of Elevated Atmospheric CO2 on Soil Fungi
--------------------------------------------------------------------------------
Reference
Rillig, M.C., Hernandez, G.Y. and Newton, P.C.D. 2000. Arbuscular mycorrhizae respond to elevated atmospheric CO2 after long-term exposure: evidence from a CO2 spring in New Zealand supports the resource balance model. Ecology Letters 3: 475-478.
What was done
The authors examined several characteristics of arbuscular mycorrhizal fungi associated with the roots of plants growing for at least 20 years along a natural CO2 gradient near a CO2-emmitting spring in New Zealand to determine the long-term effects of atmospheric CO2 enrichment on these beneficial soil fungi.

What was learned
Elevated CO2 significantly increased percent root colonization by arbuscular mycorrhizal fungi in a linear fashion - and by nearly 4-fold! - in going from 370 to 670 ppm CO2. Similarly, fungal hyphal length experienced a linear increase of over 3-fold along the same atmospheric CO2 gradient; while total soil glomalin (a protein secreted by fungal hyphae that increases soil aggregation and stability) experienced a linear increase of approximately 5-fold.

What it means
As the atmospheric CO2 concentration continues to rise, it is likely that the positive responses of arbuscular mycorrhizal fungi identified in this study will continue to become even more pronounced. If, for example, less than a doubling of the air's CO2 content has produced 3-, 4- and even 5-fold increases in fungal hyphal length, percent root colonization and total soil glomalin concentrations, respectively, what's to keep further increases in atmospheric CO2 concentration from producing 6-, 7- and 8-fold increases in these parameters? It would appear that the sky's the limit! And as these dramatic underground changes continue to occur, soil losses via wind and water erosion should be significantly reduced, due to CO2-induced glomalin-mediated increases in soil aggregate stability, which should benefit terrestrial ecosystems worldwide.

Page printed from: http://www.co2science.org/journal/2001/v4n14b2.htm Copyright © 2004. Center for the Study of Carbon Dioxide and Global Change


Rising Atmospheric CO2 Concentrations Reduce Soil Erosion: Lessons for the New Millennium
Volume 3, Number 25: 4 October 2000

--------------------------------------------------------------------------------
Over the course of his career, our father has studied a vast array of subjects related to the ongoing rise in the air's CO2 concentration; and whenever he has gained sufficient insight into a topic, he has not been afraid to state his view of its implications for the future.
One of his more prescient predictions within the context of this multifaceted subject area concerns one of earth's most valuable natural resources - its soil. Writing in his second book on the topic - Carbon Dioxide and Global Change: Earth in Transition (Idso, 1989) - our father stated that "as a result of the direct effects of atmospheric CO2 enrichment upon the primary plant processes of photosynthesis and transpiration … many plants will greatly expand their ranges with augmented water use efficiencies, stabilizing the soil and protecting it from erosion."

This prediction went against the grain of nearly all thinking on the subject at the time it was made. Starting in the 1970s and continuing almost to this day, study after study had concluded that soil erosion, via both wind and water, was running at a high sustained rate. In fact, in a recent Policy Forum article in Science, Trimble and Crosson (2000) note that "some sources have suggested that recent erosion is as great as or greater than that of the 1930s," just as some sources are suggesting that global temperatures are greater now than they were in the 1930s (Crowley, 2000; Mann 2000).

If factual, this assessment would clearly refute the prediction of our father; for with the large increase in atmospheric CO2 concentration experienced over the past 70 years, one would surely have expected to see some positive consequences, i.e., reductions in soil erosion, by now, just as one would also have expected to see significant global warming by now, if predictions of that phenomenon were correct. But herein lies the problem, or the solution to the problem, depending on one's point of view: this assessment is not factual, just as the highly-hyped global warming of the past seven decades is not factual either (see our editorials of 15 June, 1 July, and 15 July 2000).

The "remarkable feature" of the long-held belief in continued high, or even increasing, soil erosion, in the words of Trimble and Crosson, "is that it was based mostly on models," just as the global warming scare was (and still is!) based mostly on models. Indeed, they state that "little physical, field-based evidence (other than anecdotal statements) has been offered to verify the high estimates," noting that "it is questionable whether there has ever been another perceived public problem for which so much time, effort, and money were spent in light of so little scientific evidence," which almost begs us to suggest that the "perceived public problem" of CO2-induced global warming is no different, and that it will soon outstrip the soil erosion problem in this regard, if it has not already done so. But we digress; for the good news, according to Trimble and Crosson, is that "available field evidence suggests declines of soil erosion, some very precipitous, during the past six decades," which is exactly what would be expected on the basis of our father's prediction.

So what confused the issue for so many years? The problem was largely a failure to realize that most of the soil particles removed from one part of the land, by either wind or water, were deposited in nearby areas, so that the net loss of soil was only a very small portion of that which was moved about by the forces of nature.

In reviewing this shift in our perception of U.S. soil erosion history, it is interesting to note that our perceptions of several ancillary phenomena may need some adjusting too. Trimble and Crosson note, for example, that some studies have warned that "increasingly eroded soil profiles will allow less rainfall to be infiltrated and stored," leading to "increased overland flow, erosion, and flooding." But as they further note, detailed hydrologic studies indicate that just the opposite is occurring: "runoff is decreasing, flood peaks are smaller, and in some places, the base flow is greater." In addition, in their words, "these field studies show that more water is infiltrating into the soil and, in some cases, … significantly more water is being transpired by plants."

These real-world observations are also what would be expected on the basis of our father's prediction. With gradually increasing atmospheric CO2 concentrations gradually enhancing plant water use efficiencies, more plants should gradually be covering the ground, reducing rates of surface runoff and allowing more water to infiltrate into the soil, thereby providing more water to be extracted from the soil by more plants for subsequent transpiration into the air.

These hydrologic improvements, in turn, tend to improve the status of still other aspects of the planet's natural resource base, such as by increasing the stability of streams; and a good visual testament to the reality of this phenomenon is provided by a pair of photographs in the Trimble and Crosson article. Both photos show the same view of a portion of Bohemian Creek, La Crosse County, Wisconsin. The first, taken in 1940, shows an "eroded, shallow channel composed of gravel and cobbles, with coarse sediment deposited by overflows on the floodplain." The second, taken a quarter of a century later in 1974, shows that the stream channel "is narrower, smaller, and more stable." Also, "the coarse sediment has been covered with fine material, and the flood plain is vegetated to the edge of the stream." What is more, the authors note that conditions improved even more over the following 25 years.

In reviewing these many real-world manifestations of the benefits of the ongoing rise in the air's CO2 concentration for our nation's (and the world's!) important soil and water resources, we are gratified that our father had the courage to speak out on this subject as he did in his 1989 book, as well as in his earlier book (Idso, 1982). We also hope that the lesson taught by Trimble and Crosson, about the "myth and reality" of U.S. soil erosion history, will not be lost on those currently struggling with the reality of rising atmospheric CO2 concentrations and the myriad myths that have been associated with this phenomenon, such as catastrophic global warming and its attendant host of soon-to-be-experienced horror stories.

In conclusion, we agree wholeheartedly with Trimble and Crosson, who rightly state in the concluding paragraph of their important Policy Forum article that "no problem of resource or environmental management can be rationally addressed until its true space and time dimensions are known," something which has yet to be achieved in the climate change arena. And we agree even more wholeheartedly - if such is possible - with their conclusion that "the uncritical use of models is unacceptable as science and unacceptable as a basis for national policy." Unfortunately, many nations of the earth have yet to learn this lesson; and if they do not learn it soon, we could all very shortly find ourselves in a world of economic hurt, brought on by our own naivety.

Dr. Craig D. Idso
President Dr. Keith E. Idso
Vice President


References
Crowley, T.J. 2000. Causes of climate change over the past 1000 years. Science 289: 270-277.

Idso, S.B. 1982. Carbon Dioxide: Friend or Foe? An Inquiry into the Climatic and Agricultural Consequences of the Rapidly Rising CO2 Content of Earth's Atmosphere. IBR Press, Tempe, AZ.

Idso, S.B. 1989. Carbon Dioxide and Global Change: Earth in Transition. IBR Press, Tempe, AZ.

Mann, M.E. 2000. Climate change: Lessons for a new millennium. Science 289: 253-254.

Trimble, S.W. and Crosson, P. 2000. U.S. soil erosion rates - myth and reality. Science 289: 248-250.



Page printed from: http://www.co2science.org/edit/v3_edit/v3n25edit.htm
--------------------------------------------------------------------------------

Copyright © 2004. Center for the Study of Carbon Dioxide and Global Change

Monday, July 05, 2004

47. MATERIALS AND METHODS for GLOMALIN ANALYSIS 


I have included this page for serious researchers looking to do their own work. None of these appears simple or easy to me but in the right hands this is a careers worth of raw data.
GLOMALIN A Glycoproteinaceous Substance Produced by Arbuscular Mycorrhizal Fungi
MATERIALS AND METHODS for ANALYSIS
Sara Wright1 and Kristine Nichols2 1USDA-ARS Sustainable Agricultural Systems Laboratory, Beltsville, MD2Northern Great Plains Research Laboratory, Mandan, ND
INTRODUCTION: The following methods may be used to examine glomalin, an arbuscular mycorrhizal fungal protein, which is ubiquitous in the soil and has been found coating fungal hyphae and soil aggregates. Because of its importance in forming water-stable aggregates and in soil fertility, concentrations of this protein are being measured in a variety of soils to compare soils of different compositions and/or tillage or disruption practices. Please refer to the listed references for further details concerning the methodology and results. For further information on glomalin, please see the Glomalin Information page (PDF) file, Glomalin-Soil's Superglue, Glomalin: Hiding Place for a Third of the World's Stored Soil Carbon, and other Glomalin Research at Mandan, ND and Univeristy of Montana.Please e-mail Kris Nichols at nicholsk@mandan.ars.usda.gov to receive updates and changes to these procedures and/or continue to monitor the SASL Homepage and the INVAM website. Also, please let Kris know in your e-mail if you wish to be included on a list where ideas, questions, and methods may continue to be exchanged. In addition, if you have any improvements or additions to the current methods, please e-mail Kris so everyone can be informed.
TABLE OF CONTENTS
Pot culturing (PDF file)
Staining for percentage AM colonization (PDF file)
Indirect immuno-fluorescence assay (PDF file)
Glomalin extraction (PDF file)Bradford total protein assay (PDF file)Production of antibody against glomalin (PDF file)
ELISA immunoreactive protein assay (PDF file)
Immuno-dot blot assay (PDF file)
Glomalin Purification-Precipation and Dialysis (PDF file)
SDS-PAGE gel (PDF file)
Chemical and Materials Inventory (PDF file)
References (PDF file)
Disclaimer (PDF file)
For further information concerning glomalin, please see the Glomalin Information page. (PDF file)

Sunday, July 04, 2004

46. Glomalin IV Sustainability in Agriculture 

46. Glomalin IV Sustainability in Agriculture
This is an important article because it shows agricultures excited reaction to the discovery of glomalin. Agriculture fully appreciates the discovery of glomalin, and is making large scale changes in managing the soil. There are some political shots at organic farmers but the understanding of what glomalin means is right on spot. The writer also mentions the crucial role of mycorhizzia in soil health, and the restorative power of elevated carbon dioxide levels. The author can relate the Dust Bowl to ignorance in hindsight, as well as recognizing what soil with out glomalin looks like. He is also cognizant that heightened phosphorus slows or stops glomalin production, no doubt because hyphae have no need to forage for it. The article ends with a call to end plowing, and become aware of what lies beneath our feet.
Soil Discovery Endorses Conservation Tillage
By Tim Durham
GLOBAL FOOD QUARTERLY First Quarter 2003
News from the Hudson Institute’s Center for Global Food Issues
Researchers have finally found the secret of soil health and human sustainability. While critics continue to assail synthetic fertilizers and herbicides as unsustainable, a recent discovery reveals the true wildcard - a gluey soil protein that organic farmers destroy with every pass of the plow.
In 1996, Dr. Sarah Wright and colleagues at the USDA's Agricultural Research Service isolated a glycoprotein called glomalin that literally "gums up" the soil rhizosphere (the interface between soil and plant roots) with carbon fixed from the atmosphere. The compound is produced by common soil fungi called mycorrhizae that frequent the roots of many crops.
When Wright removed glomalin from soil samples, the result was a lifeless mineral powder. The soil had lost its tilth - the substance that conveys texture and health. She had inadvertently discovered the fundamental factor of soil welfare, elusive for over 10,000 years. Humic acid, previously thought to be the main contributor to soil carbon, could muster only a tiny percentage of glomalin's carbon-storing capacity in the field.
Another extraordinary finding was that elevated carbon dioxide levels encouraged mychorrizae to work overtime. Working with a consortium of scientists from UC-Davis and Stanford, Wright simulated CO2 projections for the year 2100 and observed ramped up glomalin production, with thriving fungi.
Most importantly, the USDA research demonstrated glomalin's tendency to buildup in the soil. Intensively farmed fields consistently leveled off at 0.7 mg of glomalin per gram of soil, while undisturbed plots saw an increase from 1.3 to 1.7 within three years.
In hindsight, the Dust Bowl of the 1930's wasn't a casualty of overfarming, but overplowing.
Conservation tillage maintains the supporting cast needed for soil stability, sparing mycorrhizae the stress of reestablishment every season. Aiming for at least 30% cover on the field, precision equipment gently seeds through crop residues, safeguarding soil against the elements and defending against drought.
Even before Wright's discovery, the National Soil and Water Conservation Society endorsed modern agriculture as the most sustainable in all history. According to the National Crop Residue Management Survey, 37% of corn and 57% of U.S. soybeans are now grown under some form of conservation tillage. Using herbicides and biotechnology, farmers can spray their fields with confidence, sparing produce, blighting weeds, and salvaging soil. Many more are following suit.
There are conditions, however. Members of the cabbage and spinach families are oblivious to the fungi's courtship. Growing these crops is essentially a fallow period because glomalin production stops altogether. Frequent rotation with more friendly crops is recommended.
Organic farming has two strikes against it in maintaining soil health. To satisfy nitrogen needs, crops require substantial amounts of manure. Yet manure supplies a glut of phosphorous, which shuts down glomalin production. Another complication is the near limitless supply of weed seeds bankrolled in the soil. Plowing digs up and activates seeds, causing self-induced weed outbreaks. Without herbicides, the fallback has to be the plow.
Historically, soil remains an overworked and under appreciated resource. Cornell University ecologist David W. Wolfe attributes it to pervasive "surface chauvinism", arguing that a sprinkling of soil contains countless organisms that intercept, recycle, and transform nutrients.
Mycorrhizae were noteworthy before glomalin's discovery, providing sanctuary and sustenance for a variety of soil microbes. Many of their dependents are agriculturally significant. One in particular is rhizobium, which harmlessly "infects" legumes (such as peas) to fix nitrogen from the air. Other specialized members can rapidly degrade herbicides like Roundup into carbon dioxide.
Previously labeled as an unknown and thrown away, glomalin's profound significance can only reinforce Wolfe's claim. A retiring of the plow is in order, not only to build tilth, but also to nurture the dynamic communities beneath our feet.

Glomalin III 

Glomalin III
This will wrap up my direct posting for glomalin for the time being. Serious researchers can find various ways to detect and quantify glomalin at: http://www.ba.ars.usda.gov/sasl/research/glomalin.html, Glomalin Methods and Materials. Then two more government research abstracts and another clarifier, this time by the rec.bonsai group. I have lost the link to the second article, but it is on the Tektran server.

A HYDROPHOBIC GLYCOPROTEINACEOUS SUBSTANCE PRODUCED BY MYCORRHIZAL FUNGI STABILIZES SOIL STRUCTUREAuthor(s):
MILLER R M
WRIGHT SARA E
JASTROW JULIE
UPADHYAYA ABHA

Interpretive Summary:
Sustainable agricultural practices will maintain soils as valuable natural resources. Well-aggregated soil structure is a major factor controlling the complex combination of physical, chemical, and biological processes that comprise soil ecosystem function. The discovery of a stable, insoluble, abundant glycoprotein deposited in soil by a group of ubiquitous root-associated fungi lead to the hypothesis that this protein was involved in soil aggregate stability. A chronosequence of prairie restorations was studied to determine the relationship between the protein, glomalin, and measures of aggregate size and stability. The results show that glomalin is the long-sought-after link between a healthy population of root-associated fungi (arbuscular mycorrhizal fungi) and soil aggregate stability. The goal of this work is to be able to enhance the quantity and stability of aggregates by managing the group of fungi that produce glomalin. The impact of this will be world-wide in scope and will extend to future generations.
Keywords:
soil rhizosphere ecology biology quality productivity fertility indicators transition long term sustainable agriculture study sites microbial diversity community structure biomass siderophores iron nutrient competition phospholipid fatty acid methyl ester biochemical activity characteristics
Contact:
10300 BALTIMORE BLVD.
RM 108, BG 318, BARC-EAST
BELTSVILLE
MD 20705
FAX: 301 504-8370
Email: SWRIGHT@ASRR.ARSUSDA.GOV
Approved Date: 1996-08-30

TEKTRAN
United States Department of Agriculture
Agricultural Research Service

Updated: 1998-12-18
FUNCTIONAL SIGNIFICANCE OF GLOMALIN TO SOIL FERTILITYIn the U.S., soil is lost to wind and water erosion at a rate of nearly 2 billion tons yr-1. The formation of aggregates helps stabilize soil and increase soil fertility. Organic matter concentration is correlated with the percentage of water-stable aggregates (WSA). The hypothesis of this study was that glomalin, a glycoproteinaceous substance produced by arbuscular mycorrhizal fungi, would be a major fraction of organic carbon in WSA. Four organic matter fractions – particulate organic matter (POM), total glomalin, humic acid (HA) and humin – were quantified in 1-2 mm dry-sieved aggregates from two native eastern Colorado grassland soils – Sampson and Haxtun. Each fraction, except for humin which remained in the soil after all extractions, was sequentially extracted from the same aggregate sample. Extraction procedures separated total glomalin into three different fractions: glomalin associated with POM (POM-glomalin), initial glomalin extracted from POM-free soil (glomalin), and recalcitrant glomalin (R-glomalin) extracted from POM-free soil after all other fractions. After extraction of glomalin, the POM fraction was reclassified as residual POM (R-POM). The WSA percentage was measured by wet-sieving. In this study, WSA percentage was 52% for the Sampson soil and 62% in the Haxtun soil. In the Sampson soil, the amount of carbon in the total glomalin and R-POM fractions were almost equivalent and greater than in the humin and humic acid fractions. In the Haxtun soil, the humin fraction accounted for the majority of carbon followed closely by R-POM and total glomalin. The Haxtun soil was a sandy loam with less R-POM and total glomalin, especially R-glomalin, than in the Sampson loam. The R-POM fraction contains many labile polysaccharides to help glue aggregates together as well as roots and fungal hyphae to provide the framework for aggregate formation. Glomalin also contains polysaccharides to glue aggregates together as well as iron to form stable bridges with clay minerals and hydrophobic groups such as aliphatic amino acids. The organo-mineral complexes formed between clay minerals and glomalin or humin and a hydrophobic coating from glomalin help keep aggregates water-stable and protected from water and wind erosion.

Newsgroups: rec.arts.bonsai Date: 2003-06-02 14:50:14 PST
Theo: Are you sure the article was about glucose and not glomalin? I will copy below parts of an article I published in my March 2003 newsletter. The original article was from the September 2002 issue of Agricultural Research magazine.
Until its discovery in 1996 this soil "super glue" was mistaken for an
unidentifiable constituent of soil organic matter. Rather, it permeates organic matter, binding it to silt, sand, and clay particles. Not only does glomalin contain 30 to 40 percent carbon, but it also forms clumps of soil granules called aggregates. These add structure to soil and keep other stored soil carbon from escaping.
As a glycoprotein, glomalin stores carbon in both its protein and
carbohydrate (glucose or sugar) subunits. Dr. Sarah Wright, who discovered this substance, thinks the glomalin molecule is a clump of small glycoproteins with iron and other ions attached. She found that glomalin contains from 1 to 9 percent tightly bound iron.
Glomalin is causing a complete reexamination of what makes up soil organic matter. A study showed that glomalin accounts for 27 percent of the carbon in soil and is a major component of soil organic matter. Glomalin weighs 2 to 24 times more than humic acid, a product of decaying plants that up to now was thought to be the main contributor to soil carbon. But humic acid contributes only about 8 percent of the carbon. Another team recently used carbon dating to estimate that glomalin lasts 7 to 42 years, depending on conditions.
Interestingly, the article noted that a current study in Costa Rica uses glomalin levels and root growth to measure the amount of carbon stored in soils beneath tropical forests. Researchers are finding lower levels of glomalin than expected and a much shorter lifespan. "We think it's because of the higher temperatures and moisture in tropical soils." These factors break down glomalin.(To me, this means that the extra heat our soils endure in our containerized bonsai might cause the same effects.)
It is glomalin that gives soil its tilth—a subtle texture that enables
experienced farmers and gardeners to judge great soil by feeling the smooth granules as they flow through their fingers.
Arbuscular mycorrhizal fungi, found living on plant roots around the world, appear to be the only producers of glomalin. Glomalin was named after Glomales, the taxonomic order that arbuscular mycorrhizal fungi belong to. The fungi use carbon from the plant to grow and make glomalin. In return, the fungi's hairlike filaments, called hyphae, extend the reach of plant roots. Hyphae function as pipes to funnel more water and nutrients—particularly phosphorus—to the plants.
"We've seen glomalin on the outside of the hyphae, and we believe this is how the hyphae seal themselves so they can carry water and nutrients. It may also be what gives them the rigidity they need to span the air spaces between soil particles."
As a plant grows, the fungi move down the root and form new hyphae to
colonize the growing roots. When hyphae higher up on the roots stop transporting nutrients, their protective glomalin sloughs off into the soil. There it attaches to particles of minerals (sand, silt, and clay) and organic matter, forming clumps. This type of soil structure is stable enough to resist wind and water erosion, but porous enough to let air, water, and roots move through it. It also harbors more beneficial microbes, holds more water, and helps the soil surface resist crusting.
Scientists think hyphae have a lifespan of days to weeks. The much longer lifespan of glomalin suggests that the current technique of weighing hyphae samples to estimate fungal carbon storage grossly underestimates the amount of soil carbon stored. In fact, Wright and colleagues found that glomalin contributes much more nitrogen and carbon to the soil than do hyphae or other soil microbes.
There was much more to the article, but the list limits length of responses,
so that will do for now.
Alan Walker, Lake Charles, LA, USA

===============================
Theo wrote:
HI
In the italian Issue of Bonsai and news is published a topic about
the use of glucose to enhance the roots growth. I do not know what it deals
exactly about. any feed back from others review / personal experience?
Theo

44. Glomalin II 

Glomalin II
Here are three quick items to keep the pot boiling. The first is a USDA press release. The second is Dan's post about glomalin when he saw it. I include it because of Dr. James Trappes comment about ectomycorrhiza exuding a glue. Most references to this point have concerned arbuscular mycorrhiza. My feeling is that glomalin is a structural component of hyphae and will be similar across most or all species of fungi, or mycorrhizal types at least. The author of the third article asks for quantification after a random search of the topic.
Glomalin: The Real Soil Builder
By Don Comis
February 5, 2003
An Agricultural Research Service scientist now has more proof that she has found a key ingredient responsible for the well-known benefits of soil organic matter.
Sara F. Wright, a soil scientist with the ARS Sustainable Agricultural Systems Laboratory in Beltsville, Md., discovered glomalin in 1996 and named the substance after Glomales, the taxonomic order of the fungi that produce the sticky protein. Recently, she used a nuclear magnetic resonance imager to show that glomalin is structurally different from any other organic matter component, proving it is a distinct entity.
The fungi live on most plant roots and use the plants' carbon to produce glomalin. Glomalin is thought to seal and solidify the outside of the fungi's pipelike filaments that transport water and nutrients to plants.
As the roots grow, glomalin sloughs off into the soil where it acts as a "super glue," helping sand, silt and clay particles stick to each other and to the organic matter that brings soil to life. It is glomalin that helps give good soil its feel, as smooth clumps of the glued-together particles and organic matter flow through an experienced gardener's or farmer's hands.
Glomalin was long lost in humus, the organic matter that is often called "black gold." When it did turn up in humus measurements, it was thought to be a contaminant.
Glomalin is not just the glue that holds humus to soil particles, it actually does much of what humus has been credited with. Because there is so much more glomalin in the soil than humic acid, an extractable fraction of humus, glomalin stores 27 percent of total soil carbon, compared to humic acid's eight percent. It also provides nitrogen to soil and gives it the structure needed to hold water and for proper aeration, movement of plant roots and stability to resist erosion.
ARS is the U.S. Department of Agriculture's chief scientific research agency

From: Daniel B. Wheeler (dwheeler@ipns.com)
Subject: Glomalin: fungal exudate sequesters up to a third of soil carbon for 7-42 years Newsgroups: sci.environment
Date: 2002-09-08 11:34:48 PST
Some four years after it was first published, I have found new data on
soil sequestration of carbon at:
http://www.ars.usda.gov/is/AR/archive/sep02/soil0902.htm
Glomalin: Hiding Place for a Third of World's Stored Soil Carbon
I have not had time to read all of the data yet. But it appears that
glomalin, a protein exudate produced by Glomus arbuscular mycorrhizal,
which are associated with the vast majority of plants the world over,
are also responsible for tying up significant quantities of carbon in
soil for long-term.
Glomus is further interesting to me because Glomites, the earliest
known mycorrhizal fossil known from Devonian shales dating back 400
million years, just happen to document these near-Glomus fungi which
appeared at about the same time that terrestrial plant life started to
develop.
Glomus may also be the predecessor fungi for truffles and other
ectomycorrhizal fungi, which are extremely common in recent volcanic
soils. The researcher in the above mentioned article was amazed to
find the highest levels of glomalin in Hawaii and Japan: two places
where volcanic activity is on-going. Volcanic activity along the west
coast of the US is also common, as are a plethora of ectomycorrhizal
fungi. Perhaps these ectomycorrhizal fungi also produce glomalin-like
compounds. Dr. James Trappe has noted that truffle mycelium produce
glue-like substances (similar to glomalin?) which cause soils to
absorb more water and _may_ act as root prophylactics to root rots in
PNW forests.
Daniel B. Wheeler
www.oregonwhitetruffles.com

October 31, 2003 Crumb Trail
Dirt GlueAn unanticipated benefit to blogging for me comes from examining access logs for search strings used by others to find the Crumb Trail. Some are humorous or salacious but others are useful. I sometimes google interesting strings myself. Today I googled "glomalin methods". Crumb Trail was 40 or so links down the list so whoever did the search worked to find it. At the top of the list was a USDA page devoted to sharing techniques for measuring glomalin.
The following methods may be used to examine glomalin, an arbuscular mycorrhizal fungal protein, which is ubiquitous in the soil and has been found coating fungal hyphae and soil aggregates. Because of its importance in forming water-stable aggregates and in soil fertility, concentrations of this protein are being measured in a variety of soils to compare soils of different compositions and/or tillage or disruption practices. Please refer to the listed references for further details concerning the methodology and results.
There was also a link to a glomalin information page
Soil aggregation is a complex process that is largely dependent upon microorganisms to provide glues that hold soil particles together. These glues are carbon-containing compounds that protect microorganisms from drying out. We are beginning to understand the importance of one group of soil fungi and the glue that is produced in large amounts by these fungi. The fungi are the arbuscular mycorrhizal fungi (AMF) and the glue was named glomalin after Glomales — the taxonomic order of this group of fungi..
Glomalin concentration and aggregate stability are related over 3 years during conversion from conventional tillage (P-T) to no tillage (N-T) corn (Fig. 3). A comparison was made with a perennial grass that grew undisturbed for 15 years as a buffer around the plots. Increases in both stability and glomalin were seen at yearly intervals, but had not reached the levels in the undisturbed grass. Higher levels of glomalin give greater water infiltration, more permeability to air, better root development, higher microbial activity, resistance to surface sealing (crusts) and erosion (wind/water).
Marvelous stuff. If we stop plowing fields will be more fertile with more organic matter, retain moisture as well as drain well and make better use of phosphorous and so need less fertilizers. Soil life will increase and plants will thrive.
Croplands will never be as healthy as grasslands but we can do much better. It is worth noting that soils get better year after year when left untilled as glomalin increases. It might be interesting to attempt to quantify the amount of carbon that can be sequestered this way since glomalin is so durable, accumulating year after year rather than cycling back to the atmosphere.












Saturday, July 03, 2004

43. Glomalin I 

43. Glomalin I
This is the article I first found in regard to glomalin. After it are two abstracts that contain suggestions for future research. There is also an earlier press release and a page of sampling methods for researchers I will post.


Glomalin: Hiding Place for a Third of the World's Stored Soil Carbon A sticky protein seems to be the unsung hero of soil carbon storage.


Until its discovery in 1996 by ARS soil scientist Sara F. Wright, this soil "super glue" was mistaken for an unidentifiable constituent of soil organic matter. Rather, it permeates organic matter, binding it to silt, sand, and clay particles. Not only does glomalin contain 30 to 40 percent carbon, but it also forms clumps of soil granules called aggregates. These add structure to soil and keep other stored soil carbon from escaping.


As a glycoprotein, glomalin stores carbon in both its protein and carbohydrate (glucose or sugar) subunits. Wright, who is with the Sustainable Agricultural Systems Laboratory in Beltsville, Maryland, thinks the glomalin molecule is a clump of small glycoproteins with iron and other ions attached. She found that glomalin contains from 1 to 9 percent tightly bound iron.

Glomalin is causing a complete reexamination of what makes up soil organic matter. It is increasingly being included in studies of carbon storage and soil quality. In fact, the U.S. Department of Energy, as part of its interest in carbon storage as an offset to rising atmospheric carbon dioxide (CO2) levels, partially funded a recent study by lab technician Kristine A. Nichols, a colleague of Wright's. Nichols reported on the study as part of her doctoral dissertation in soil science at the University of Maryland.

That study showed that glomalin accounts for 27 percent of the carbon in soil and is a major component of soil organic matter. Nichols, Wright, and E. Kudjo Dzantor, a soil scientist at the University of Maryland-College Park, found that glomalin weighs 2 to 24 times more than humic acid, a product of decaying plants that up to now was thought to be the main contributor to soil carbon. But humic acid contributes only about 8 percent of the carbon. Another team recently used carbon dating to estimate that glomalin lasts 7 to 42 years, depending on conditions.

For the study, the scientists compared different chemical extraction techniques using eight different soils from Colorado, Georgia, Maryland, and Nebraska. They found that current assays greatly underestimate the amount of glomalin present in soils. By comparing weights of extracted organic matter fractions (glomalin, humic acid, fulvic acid, and particulate organic matter), Nichols found four times more glomalin than humic acid. She also found that the extraction method she and Wright use underestimates glomalin in certain soils where it is more tightly bound than usual.


In a companion study, Nichols, Wright, and Dzantor teamed up with ARS chemist Walter F. Schmidt to examine organic matter extracted from the same soils under a nuclear magnetic resonance (NMR) imager. They found that glomalin's structure differs from that of humic acid—or any other organic matter component—and has unique structural units.

In a current study in Costa Rica, partly funded by the National Science Foundation, Wright is using glomalin levels and root growth to measure the amount of carbon stored in soils beneath tropical forests. She is finding lower levels of glomalin than expected and a much shorter lifespan. "We think it's because of the higher temperatures and moisture in tropical soils," she explains. These factors break down glomalin.


Forests, croplands, and grasslands around the world are thought to be valuable for offsetting carbon dioxide emissions from industry and vehicles. In fact, some private markets have already started offering carbon credits for sale by owners of such land. Industry could buy the credits as offsets for their emissions. The expectation is that these credits would be traded just as pollution credits are currently traded worldwide.

How Does Glomalin Work?


It is glomalin that gives soil its tilth—a subtle texture that enables experienced farmers and gardeners to judge great soil by feeling the smooth granules as they flow through their fingers.


Arbuscular mycorrhizal fungi, found living on plant roots around the world, appear to be the only producers of glomalin. Wright named glomalin after Glomales, the taxonomic order that arbuscular mycorrhizal fungi belong to. The fungi use carbon from the plant to grow and make glomalin. In return, the fungi's hairlike filaments, called hyphae, extend the reach of plant roots. Hyphae function as pipes to funnel more water and nutrients—particularly phosphorus—to the plants.


"We've seen glomalin on the outside of the hyphae, and we believe this is how the hyphae seal themselves so they can carry water and nutrients. It may also be what gives them the rigidity they need to span the air spaces between soil particles," says Wright.

As a plant grows, the fungi move down the root and form new hyphae to colonize the growing roots. When hyphae higher up on the roots stop transporting nutrients, their protective glomalin sloughs off into the soil. There it attaches to particles of minerals (sand, silt, and clay) and organic matter, forming clumps. This type of soil structure is stable enough to resist wind and water erosion, but porous enough to let air, water, and roots move through it. It also harbors more beneficial microbes, holds more water, and helps the soil surface resist crusting.


Scientists think hyphae have a lifespan of days to weeks. The much longer lifespan of glomalin suggests that the current technique of weighing hyphae samples to estimate fungal carbon storage grossly underestimates the amount of soil carbon stored. In fact, Wright and colleagues found that glomalin contributes much more nitrogen and carbon to the soil than do hyphae or other soil microbes.


Rising CO2 Boosts Glomalin, Too


In an earlier study, Wright and scientists from the University of California at Riverside and Stanford University showed that higher CO2 levels in the atmosphere stimulate the fungi to produce more glomalin.


They did a 3-year study on semiarid shrub land and a 6-year study on grasslands in San Diego County, California, using outdoor chambers with controlled CO2 levels. When CO2 reached 670 parts per million (ppm)—the level predicted by mid to late century—hyphae grew three times as long and produced five times as much glomalin as fungi on plants growing with today's ambient level of 370 ppm.


Longer hyphae help plants reach more water and nutrients, which could help plants face drought in a warmer climate. The increase in glomalin production helps soil build defenses against degradation and erosion and boosts its productivity.


Wright says all these benefits can also come from good tillage and soil management techniques, instead of from higher atmospheric CO2.


"You're in the driver's seat when you use techniques proven to do the same thing as the higher CO2 that might be causing global warming. You can still raise glomalin levels, improve soil structure, and increase carbon storage without the risks of the unknowns in global climate change," she says.

Putting Glomalin to Work


Wright found that glomalin is very manageable. She is studying glomalin levels under different farming and ranching practices. Levels were maintained or raised by no-till, cover crops, reduced phosphorus inputs, and the sparing use of crops that don't have arbuscular mycorrhizal fungi on their roots. Those include members of the Brassicaceae family, like cabbage and cauliflower, and the mustard family, like canola and crambe.


"When you grow those crops, it's like a fallow period, because glomalin production stops," says Wright. "You need to rotate them with crops that have glomalin-producing fungi."


In a 4-year study at the Henry A. Wallace Beltsville (Maryland) Agricultural Research Center, Wright found that glomalin levels rose each year after no-till was started. No-till refers to a modern conservation practice that uses equipment to plant seeds with no prior plowing. This practice was developed to protect soil from erosion by keeping fields covered with crop residue.


Glomalin went from 1.3 milligrams per gram of soil (mg/g) after the first year to 1.7 mg/g after the third. A nearby field that was plowed and planted each year had only 0.7 mg/g. In comparison, the soil under a 15-year-old buffer strip of grass had 2.7 mg/g.


Wright found glomalin levels up to 15 mg/g elsewhere in the Mid-Atlantic region. But she found the highest levels—more than 100 mg/g—in Hawaiian soils, with Japanese soils a close second. "We don't know why we found the highest levels in Hawaii's tropical soils. We usually find lower levels in other tropical areas, because it breaks down faster at higher temperature and moisture levels," Wright says. "We can only guess that the Hawaiian soils lack some organism that is breaking down glomalin in other tropical soils—or that high soil levels of iron are protecting glomalin."

It's Persistent and It's Everywhere!


The toughness of the molecule was one of the things that struck Wright most in her discovery of glomalin. She says it's the reason glomalin eluded scientific detection for so long.


"It requires an unusual effort to dislodge glomalin for study: a bath in citrate combined with heating at 250 °F for at least an hour," Wright says. "No other soil glue found to date required anything as drastic as this."


"We've learned that the sodium hydroxide used to separate out humic acid in soil misses most of the glomalin. So, most of it was thrown away with the insoluble humus and minerals in soil," she says. "The little bit of glomalin left in the humic acid was thought to be nothing more than unknown foreign substances that contaminated the experiments."


Once Wright found a way to capture glomalin, her next big surprise was how much of it there was in some soils and how widespread it was. She tested samples of soils from around the world and found glomalin in all.


"Anything present in these amounts has to be considered in any studies of plant-soil interactions," Wright says. "There may be implications beyond the carbon storage and soil quality issues—such as whether the large amounts of iron in glomalin mean that it could be protecting plants from pathogens."


Her recent work with Nichols has shown that glomalin levels are even higher in some soils than previously estimated.


"Glomalin is unique among soil components for its strength and stability," Wright says. Other soil components that contain carbon and nitrogen, as glomalin does, don't last very long. Microbes quickly break them down into byproducts. And proteins from plants are degraded very quickly in soil.


"We need to learn a lot more about this molecule, though, if we are to manage glomalin wisely. Our next step is to identify the chemical makeup of each of its parts, including the protein core, the sugar carbohydrates, and the attached iron and other possible ions." Nichols is starting to work on just that.


"Once we know what sugars and proteins are there," says Nichols, we will use NMR and other techniques to create a three-dimensional image of the molecule. We can then find the most likely sites to look for iron or other attached ions.


"Researchers have studied organic matter for a long time and know its benefits to soil. But we're just starting to learn which components of organic matter are responsible for these benefits. That's the exciting part of glomalin research. We've found a major component that we think definitely has a strong role in the benefits attributed to organic matter—things like soil stability, nutrient accessibility, and nutrient cycling."


As carbon gets assigned a dollar value in a carbon commodity market, it may give literal meaning to the expression that good soil is black gold. And glomalin could be viewed as its golden seal.—By Don Comis, Agricultural Research Service Information Staff.


This research is part of Soil Resource Management, an ARS National Program (#202) described on the World Wide Web at http://www.nps.ars.usda.gov/.


Sara F. Wright and Kristine A. Nichols are with the USDA-ARS Sustainable Agricultural Systems Laboratory, Bldg. 001, 10300 Baltimore Ave., Beltsville, MD 20705; phone (301) 504-8156 [Wright], (301) 504-6977 [Nichols], fax (301) 504-8370.

"Glomalin: Hiding Place for a Third of the World's Stored Soil Carbon" was published in the September 2002 issue of Agricultural Research magazine.





SOIL ORGANIC MATTER COMPONENTS II: CHARACTERIZATION OF GLOMALIN AS A UNIQUE COMPONENT OF SOIL ORGANIC MATTERAuthor(s):

NICHOLS KRISTINE A
WRIGHT SARA E
SCHMIDT WALTER F
DZANTOR E K


Interpretive Summary:

Interventions to maintain soil productivity and sustainability depend upon understanding soil biology. An important biologically-derived component that contributes to soil health is organic matter. Currently, the organic matter constituents that are studied to determine sustainable agricultural practices are humic acid (HA), fulvic acid (FA and particulate organic matter (POM). In 1996, we discovered an abundant glycoprotein, glomalin, in soil. Glomalin is produced by a ubiquitous root-colonizing group of fungi - the arbuscular mycorrhizal fungi. It was important to determine why glomalin had not been found by very competent scientists during 50 years of research on HA and FA and 10 years of research on POM. Chemical co-extraction of HA and FA was performed, and then the extraction of glomalin on the residual soil. Also, glomalin was extracted first , and then HA and FA were co-extracted from the residual soil. This study showed that the methods used to extract HA and FA are not effective in extracting even small amounts of glomalin. Also, POM contains glomalin but chemical extractions were not previously used to show components of POM. This studied showed that glomalin was unique in composition and structure in comparison to HA, FA and POM. This information will be valuable to scientist and farmers throughout the world. Agricultural practices that encourage proliferation of the fungi that produce glomalin can be defined to prevent soil degradation, and degraded soil can be brought back to productivity.


Keywords:

microbial methods agroecosystems beneficial microorganisms glomalin mycorrhiza microbial ecology

TEKTRAN

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SOIL MICROBIAL BIOMASS AND MINERALIZABLE CARBON OF WATER-STABLE AGGREGATES AFFECTED BY TEXTURE AND TILLAGE
Author(s):

FRANZLUEBBERS ALAN J
ARSHAD M A


Interpretive Summary:

Knowledge of soil organic carbon sequestration and turnover is important in understanding biogeochemical cycles, land management effects on soil quality, and the contribution of soils to greenhouse gas emissions. Macroaggregates (>0.25 mm diam.) had a greater concentration of soil microbial biomass carbon and mineralizable carbon than microaggregates near the soil surface, but more similar concentrations of these active carbon pools at lower depths. We observed a redistribution of active carbon pools within water-stable aggregate classes suggesting that active carbon pools became more associated with macroaggregates under zero tillage and with microaggregates under conventional tillage. Soil under zero tillage contained more macroaggregate-protected organic carbon than under conventional tillage, indicating that reduced soil disturbance could lead to greater organic carbon sequestration and improved soil quality with improvements in aggregation.


Keywords:

agroecosystems biogeochemical watersheds nutrients rootzone


Contact:

SOU. PIEDMONT CONS. RES.
1420 EXPERIMENT STA ROAD
WATKINSVILLE
GA 30677-2373
FAX: 706/769-8962
Email: afranz@uga.cc.uga.edu


Approved Date: 1996-08-22


--------------------------------------------------------------------------------

TEKTRAN
United States Department of Agriculture
Agricultural Research Service

Updated: 1998-12-18



Contact:

10300 BALTIMORE AVE.
BLDG 001, ROOM 342, BARC-
BELTSVILLE
MD 20705
FAX: (301)504-6491
Email: wrights@ba.ars.usda.gov


Approved Date: 2001-12-27

42. CO2 Fertilization May Be Slowing Global Warming 

This is a brand new study that supports CO2 Science claims that elevated CO2 will be absorbed by biological systems. Knowing this we can harness it into forms we want or need, like reforestation. The release of CO2 from soils as dust is interesting to see included. We are glad to see forest studies appearing, especially in connection with climate science. Still, the need to quantify glomalin and carbon storage in forest systems is only emerging. The role of CO2 in gluing landscapes together hasn't been investigated, although I am sure someone is starting to wonder why revegetation should restore devastated ecosytems so well.
Source: Boston College
Date: 2004-06-18
URL: http://www.sciencedaily.com/releases/2004/06/040618065651.htm
CO2 Fertilization May Be Slowing Global Warming
CHESTNUT HILL, MA (6-17-04) --- A Boston College scientist has published new research introducing the concept of a CO2 fertilization factor for soil carbon, a way to measure an ecosystem's ability to store carbon in response to increased carbon dioxide in the atmosphere. The study, authored by Kevin G. Harrison, an assistant professor in Boston College's Geology and Geophysics Department, has serious implications for scientists examining global climate change who have long sought information on missing carbon sinks.

His research appears in the May 2004 Geochemistry, Geophysics, Geosystems (G3), an electronic journal published by the American Geophysical Union, which showcases discoveries in geophysics and geochemistry that cross traditional disciplinary boundaries and approach the Earth as a system.

Harrison's research says that CO2 fertilization may be slowing down the expected accumulation of carbon dioxide in the atmosphere by increasing carbon accumulation in terrestrial vegetation and soil.

"I have determined a CO2 fertilization factor of 1.18 for a white oak ecosystem using soil carbon and radiocarbon measurements. If major terrestrial ecosystems have similar values, CO2 fertilization may be transferring enough carbon from the atmosphere to the soil to balance the global carbon budget," according to Harrison.

"It is my hope that these concepts will be used by global change geochemists worldwide," writes Harrison. Samples for the study were collected from a white oak experiment at the Global Change Field Research Site in Oak Ridge, Tenn. The research has been funded by the U.S. Department of Agriculture.

Harrison's research focuses on the effects of fossil fuel combustion, dust and deforestation on the buildup of carbon dioxide in the atmosphere. At Boston College, he teaches courses on "Biogeochemistry of the Habitable Planet"; "Environmental Geochemistry: Living Dangerously," and "Global Warming." He earned a bachelor of science degree in chemistry at Brown University. He received a master's degree in marine chemistry from the University of California at San Diego's Scripps Institution of Oceanography, and master's and doctoral degrees in geological sciences from Columbia University.


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This story has been adapted from a news release issued by Boston College.

41.Fungi (In Association with Woody Plants)-- Summary 

I continue to lay out my primary sources for those who care to read along and do their own research and/or draw their own conclusions. Here we can see elevated carbon dioxide is accelerating aboveground and total biomass, root tip colonization by fungi,hyphae growth, and glomalin production.

Fungi (In Association with Woody Plants) -- Summary
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As the CO2 content of the air increases, nearly all of earth's plants will exhibit increases in their photosynthetic rates. Additional photosynthetic sugars can then be utilized to increase plant growth and development, or they can be exuded belowground to enhance the growth and development of symbiotic fungal organisms that live in close association with plant roots. This latter phenomenon is extremely important, for fungi form intricate mycelial networks that act to increase nutrient and water harvesting from the soil and the subsequent transfer of these important resources to plants. In this summary, we review how elevated CO2 impacts these several processes in the case of woody species, i.e., trees and shrubs.
Researchers often report that atmospheric CO2 enrichment increases the percentage of the root tip that is colonized by the hyphae of ectomycorrhizal fungi (Staddon and Fitter, 1998). Rygiewicz et al. (2000), for example, observed a 95% increase in this parameter for Douglas fir seedlings exposed to an atmospheric CO2 concentration of 550 ppm for four years. Similarly, Godbold et al. (1997) noted that the percentage root tip colonized by ectomycorrhizal fungal hyphae in paper birch and Eastern white pine were enhanced by 13 and 38%, respectively, when the atmospheric CO2 concentration was doubled for nearly ten months. In addition, other scientists have documented this positive influence of elevated CO2 on the percentage root tip colonized by fungal hyphae in yellow birch (+71%, Berntson and Bazzaz, 1998), white birch (+45%, Berntson and Bazzaz, 1998), and Eastern hemlock (+47%, Godbold et al., 1997).

Although the percentage increase in root colonization by fungal hyphae is by far the most commonly reported response of fungi to atmospheric CO2 enrichment, it is not the only way in which elevated levels of atmospheric CO2 enhanced fungal-root relationships.. In the study of Rillig and Allen (1998), for example, elevated CO2 (750 ppm) did not significantly impact the percentage root colonized by fungal hyphae in Gutierrezia sarothrae shrubs, but it did significantly increase its percentage colonization by arbuscules and vesicles by 14- and 2.5-fold, respectively. Similarly, Klironomos et al. (1998) reported that twice-ambient levels of atmospheric CO2 increased the percentage root colonization in sagebrush by arbuscules by 70%. In addition, Rillig et al. (2000) documented a 4-fold linear increase in the percentage root colonization by arbuscular mycorrhizal fungi in native vegetation growing for 20 years along a natural CO2 gradient (370 to 670 ppm) near a CO2 spring in New Zealand.

Elevated CO2 has also been noted to significantly enhance other fungal properties. In the study of Walker et al. (1998), an atmospheric CO2 concentration of 700 ppm increased the total number of ectomycorrhizal hyphae on ponderosa pine seedling roots by 85% relative to what was observed on roots of seedlings exposed to ambient air. In the more complex study of Wiemken et al. (2001), the biomass of ectomycorrhizal fungi associated with roots of beech and spruce seedlings was about 118% greater at high soil nitrogen and elevated CO2 (560 ppm) than it was at low soil nitrogen and ambient CO2.

Finally, atmospheric CO2 enrichment also impacts certain fungal properties that can act to enhance soil stability. In the previously mentioned study of Rillig et al. (2000), for example, it was reported that total soil glomalin - a protein secreted by fungal hyphae that increases soil aggregation and stability - increased nearly 5-fold along the natural 300-ppm CO2 gradient. In addition, Rouhier and Reed (1998) noted that the mycorrhizal network constructed around roots of Scots pine seedlings grown at 700 ppm CO2 occupied 444% more soil area than that occupied by networks constructed around roots of seedlings exposed to ambient air. Rouhier and Read (1999) documented the same phenomenon for mycorrhizal networks established around roots of CO2-enriched beech seedlings, albeit the response was much smaller in magnitude (30% increase).

In summary, as the air's CO2 content continues to rise, most of earth's woody species should respond by exhibiting enhanced rates of photosynthesis that will lead to the stimulation of belowground fungal development (increased fungal hyphal length and biomass) and activities (greater penetration of the soil, enhanced soil stability), leading ultimately to enhanced root uptake of important soil minerals and water.

References
Berntson, G.M. and Bazzaz, F.A. 1998. Regenerating temperate forest mesocosms in elevated CO2: belowground growth and nitrogen cycling. Oecologia 113: 115-125.

Godbold, D.L., Berntson, G.M. and Bazzaz, F.A. 1997. Growth and mycorrhizal colonization of three North American tress species under elevated atmospheric CO2. New Phytologist 137: 433-440.

Klironomos, J.N., Ursic, M., Rillig, M. and Allen, M.F. 1998. Interspecific differences in the response of arbuscular mycorrhizal fungi to Artemisia tridentata grown under elevated atmospheric CO2. New Phytologist 138: 599-605.

Rillig, M.C. and Allen, M.F. 1998. Arbuscular mycorrhizae of Gutierrezia sarothrae and elevated carbon dioxide: evidence for shifts in C allocation to and within the mycobiont. Soil Biology and Biochemistry 30: 2001-2008.

Rillig, M.C., Hernandez, G.Y. and Newton, P.C.D. 2000. Arbuscular mycorrhizae respond to elevated atmospheric CO2 after long-term exposure: evidence from a CO2 spring in New Zealand supports the resource balance model. Ecology Letters 3: 475-478.

Rouhier, H. and Read, D.J. 1998. Plant and fungal responses to elevated atmospheric carbon dioxide in mycorrhizal seedlings of Pinus sylvestris. Environmental and Experimental Botany 40: 237-246.

Rouhier, H. and Read, D. 1999. Plant and fungal responses to elevated atmospheric CO2 in mycorrhizal seedlings of Betula pendula. Environmental and Experimental Botany 42: 231-241.

Rygiewicz, P.T., Martin, K.J. and Tuininga, A.R. 2000. Morphotype community structure of ectomycorrhizas on Douglas fir (Pseudotsuga menziesii Mirb. Franco) seedlings grown under elevated atmospheric CO2 and temperature. Oecologia 124: 299-308.

Staddon, P.L. and Fitter, A.H. 1998. Does elevated atmospheric carbon dioxide affect arbuscular mycorrhizas? Trends in Ecology and Evolution 13: 455-458.

Walker, R.F., Johnson, D.W., Geisinger, D.R. and Ball, J.T. 1998. Growth and ectomycorrhizal colonization of ponderosa pine seedlings supplied different levels of atmospheric CO2 and soil N and P. Forest Ecology and Management 109: 9-20.

Wiemken, V., Ineichen, K. and Boller, T. 2001. Development of ectomycorrhizas in model beech-spruce ecosystems on siliceous and calcareous soil: a 4-year experiment with atmospheric CO2 enrichment and nitrogen fertilization. Plant and Soil 234: 99-108.



Page printed from: http://www.co2science.org/subject/f/summaries/fungiwood.htm
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Copyright © 2004. Center for the Study of Carbon Dioxide and Global Change

Friday, July 02, 2004

40. Restoration of a native plant system. 

40. Restoration of a native plant system.
From: Manual of California Native Plants
This article has really useful information regarding restoration with native species of plants, why and why nots and dos and don’ts. He points out the mycorhizzial ground between plants, probable duration of restarting fungal activity, weeds and grasses, cattle and how to choose a native plant nurseryman. My main point here is the entire ground is part of the network and is glomalin conditioned to hold water, and that native plantsthrive when allowed because of mycological associations. Another great round of opportunities for research and plant science work, and we concur with the lack of available data on conditions over the last 500 years. Click on the link to Las Pilatas Nursery for an outstanding collection of over 1500 text pages dealing with California Native plants, ecosystems, wildlife,etc.
Restoration of a native plant system (Putting your ecology back on track.)
In our Calif. ecosystems the distant past(greater than 10,000 years) has little or no relevance on the present. We are trying to find historical accounts of the period in the 1700's when the plant communities were last stable. After that, fire frequency changes. The records even in the 1800's are bad at best. Our ecology is changing so fast we are having trouble documenting plant assoc. on a site (never mind mycorrhizal assoc.) from only 20-30 years before. Our site is a case in point. The neighbors that have moved in since the 1985 fire assume the hillsides always looked as they do now. Only the largest crown sprouters remain the same.
We have been trying to revegetate back to the original communities using every trick we can think of. So far we have found:
1. No site is special or mystical. All react appropriately if you include past plant community, (ideally would be to track the last 500 years) mycorrhizal assoc., soil and climate in your plan of attack.
2. You can only successfully revegetate plant species that are members of the original plant community or close to it. The grassland community will best support the grassland species and the chaparral community will best support the chaparral species. Trying to make an area other than what it is will bring in the pioneers from the appropriate community. This is why history is important. As an example, Baccharis moves into the Coastal Sage Scrub areas that have been over the years converted into grasslands. The plant community reverts to the original plant community, as we stated earlier. It may take 200 years but if left alone it generally will. The roots and mycorrhizal assoc. will move about 2-5 foot per year. If a colonizer can become established a herbivore will usually carry a fungal inoculum to it.
We've been told the prairie was easy to restore, 'just burn it off after 3-5 years, the non-natives cannot take the fire'. That was in the mid-west prairie with their protocols. That would be very detrimental to our Calif. ecosystems! Our communities appear to be much longer term because of moisture stress, and possess a dual cycle with perennials, shrubs and trees. Again, each community needs to be examined on its own, each climate is different and each soil type is different. The coastal prairie is different from the valley grassland and much different from the mid-west prairie or the African savanna by climates, soils, and plant species.
3. If the plant community has any vitality left in it you can use it to help you by planting community-supported plants and letting the strengthened community help you suppress the non-native weeds. We have been planting one-watered native landscaping for two years now using this approach. We only water once, forever. You have to follow carefully the plant community, climate and soil of each site for this to work. But it has worked to date in the following communities; Chaparral, Coastal Sage Scrub, Foothill Woodland, Closed Cone Pine Forest, Coastal Prairie, and Yellow Pine forest. We have had failure only in the riparian community.
To determine if the mycorrhizal grid is still intact. Are there any native species left on the site? If so, is there intact(non-cut with rippers and ditches) soil between plants? If so, the areas between plants can be considered mycorrhizal. Has only the top few inches of the soil been disturbed even though the surface plants are removed? If the disturbance was within the last 4 years or so, (a few systems last longer, most are viable for 2-5 years) you can treat it as a mycorrhizal site and restart the system.
The weed suppression is dramatic in many undisturbed communities on many weeds (through allelopathy, gradual buildup of the litter layer, the fact that most of the mycorrhizae that help each plant in the community to survive through increased availability of nutrients and suppression of pathogens are not in association with the weedy, introduced species and all plants and animals in the community from microorganisms up actively attack any non-community specific species up to a certain point. An analogy would be that the suppression of weeds by an undisturbed plant community is like suppression of disease by our body's immune system). You'll find a few survive long term but not many. Next time you go into an area with disturbance next to a pristine area look how the weeds are having trouble extending into the undisturbed area. With a little help from you and I they cannot. This includes grasslands.
3a. You have to suppress the growth of the weeds or remove the weeds at the correct time of year that is appropriate for the community upon which you are working, with minimum disturbance. Any disturbance of the soil or addition of water or fertilizer is detrimental to the native plants but encourages the growth of the weeds(ruderals). Timing varies by community, climate, soil, and sometimes by species. That is one of the reasons we look at sprays on many sites. Soil disturbance of any type, no matter how well intentioned is still an infection point. But you need to know your site's plants before you commit to spray. We know of no other way to remove the non-natives and ruderals without soil disturbance in a short period of time, if we are trying to recover or protect a community in years instead of centuries. Whether it be by tractor disc or human hand it is still soil disturbance and on bad sites this is not acceptable.
4. Knowing that the weeds emerge before the native plants has caused us to try spraying with Round-up to suppress the weeds and encourage the later emerging natives. (Trappe et. al. lists Round-up as no problem for mycorrhiza and our experience has been it kills only what is green with little or no inhibition of later seedlings.) This has proven to be somewhat effective as most (but not all) perennial grasses and dec. shrubs are brown at this time (early Dec.) and not bothered by the sprays. This has to be done very early or very carefully though because the bulbs and native annuals come up in at this same time along with some of the perennial grasses sprouting about then. A very short term pre-emergent may be the answer although many of the ruderals are resistant to pre-emergents and early native sprouters will be hurt. You really need to do a site survey year to year before to see if there are rare plants on site and if the site is evolving native or non-native. Can you make a spray in a non-lethal way to your on-site natives? We've found this early approach to be most effective if used during drought years after the site's energy has been recharged. Do not use it during wet years unless you follow up with a January community-specific wildflower seeding. That is the only way you'll see wildflowers in many of the disturbed communities. Spraying is only an alternative when plant communities have been almost destroyed and need to be helped to recover their balance short-term. The tradeoff, do we lose a few common species to save the long cycle fire-followers?
5. Knowing that another set of the non-natives is usually green and set seed after the wildflowers set seed led us to try spot spraying with Round-up afterwards. You also have to cover each perennial before you spray. This proved to be more effective but the timing has to be exact. The mustard and Starthistle have to be in mid-flower when they have committed their energy to reproduction but before they have set seed. Again, it should be sprayed in drought years if possible.
6a. We tried grazing down the ruderals but the horses selectively ate the native bunch grasses and annual grasses further tweaking the disturbed community in favor of the ruderals. We've seen the same thing happen with cattle on other sites(they ate the perennials and trees also). We also noticed that the weeds that were in the hay but not in the ryegrass were very few indeed. These radiated away from the horse's feeding pen. Although this secondary source was relatively minor it should be watched as a problem in strict sites. (Along horse paths.)
6b. We tried mowing down the weeds to create a litter layer but the weeds can set seed when they are only an inch tall!
7a. From these tries over a 5-year program we offer these theories on communities with seasonal wildflowers.
7b. The displacement of weeds(ruderals) and non-natives is possible in most of our native ecosystems as long as some of the plant community is intact.
7c. No single approach will work. All levels of the community from the microorganisms up must be helped.
7d. If two sprayings, early and late are used, you must use plant community specific pioneer wildflowers to cover the ground and charge the grid during the critical spring 'energy capture'. Covering the ground is important as the weeds have trouble infecting a closed biosystem and covered ground. We are in favor of using species that are widely in use, native throughout most of Calif. and do not need to be site specific. There have been a series of almost criminal substitutions for narrow site specific plants. (Non-native Bromus for the native one. Sometimes a sub-species from S.Cal. gets used in N.CAL. and such, ruining the sub-species.) We would rather see the same species used every time state wide so we at least know where the aliens came from. Our inclination is to divide the poppies into Munz's (1974) 4 sub-forms of Eschscholzia and use the appropriate one in the right area of the state, combined with Lupinus succulentus, Lupinus polycarpus, Lupinus bicolor ssp. marginatus, and Clarkia purpurea ssp. quad..
Most of the pretty, showy wildflowers are either pioneer plants in plant communities where they get buried with vegetation or an important part of the long term ecosystem in grasslands, deserts and prairies. If you are planting wildflowers in communities other than prairie, grassland, meadow, foothill woodland, and most desert sites your wildflowers will be unstable pioneers and disappear after a few years.
Mother nature does this after a fire with fire following pioneer plants covering the ground, feeding the mycorrhizal grid, and then declining and dying, further feeding the grid as the climax seedlings mature in.
7e. We feel grasses cannot be replanted back into many of these sites until you have at least a partial control of weeds as it limits your strategies. Planting the larger summer species would be a better alternative, (for example the right shrub Buckwheat (Eriogonum sp.) for the site.)
7f. It may take 5-70 years of active restoration strategies to achieve a stable, viable, native plant community.
7g. We're realists and realize that although we think they are an ecological disaster, cattle will not go away. Grazers can be used in place of Round-up but only within windows that allow the wildflowers to grow and seed with no stresses and the perennials to set seed during the wet years. In much of Calif. this would mean that cattle are only acceptable from July through Dec., and a second period in May-June during the extreme drought years. During years of normal or higher rainfall, keep cattle numbers down. There are very serious down sides to cattle. They weigh too much. Compacted soil suppresses mycorrhiza and encourages weeds. They tear holes in the light litter that covers many native communities giving entry points for the always aggressive weeds. They carry non-native seeds from their past(previous pasture) and infect everywhere they go. They also favor the worst weeds over the natives by eating the more palatable natives first.
7h. You must work with community, climate and soil-specific ideas. You must work with the strategies of that community to help that community fend off the invading non-native species.
7i. The community can only move out and recapture an area a few feet a year, usually with stops and starts. Start in one area and work out. (See the Bradley Method..Fremontia, Vol.13, No.2)
7j. Our native community works on a beneficial cycle and the introduced weeds disrupt this cycle. This shows up very well in areas that are disturbed next to an intact community. The weeds try to move out into the community but are strongly suppressed as long as the community is intact. If the fungal-plant community fails, the weeds take over. A site covered with weeds in the middle of a plant community is indicative of a struggling community.
7k. If you need to reseed for some reason, each site needs to be treated with some thought. Most intact sites should not be reseeded. We usually use the correct poppy and lupine for the site as the pioneer species to re-hab within a VAM (vesicular-arbuscular mycorrhiza) system. These facultative mycotrophs have proven to be what rye grass was billed to be, benign to the environment but supportive of the community and soil structure.
If the site is mid to low elevation Ectomycorrhizal (pine forests, oak woodland, mixed evergreen) use a combination of lupines, deerweed(Lotus), and native vetch to reseed if needed. No grass. Our preferred plants are lupines as they are facultatively VAM nitrogen fixers and have roots that are among the strongest soil penetrators. Higher elevation forest sites are best left alone because community specific seed is not available.
Finally, the best way to preserve rare plants is to not graze, seed or disturb a site. Check your county, most are still recommending seeding all disturbed sites with ryegrass. Send them a copy of this article, hopefully you will save an ecosystem.
Contract Grown, Site Specific plant material
Many sites, from a single homeowner, to a large pipeline need site specific plant material grown.
Much of what we grow goes to these sites. We are hearing of, and seeing requirements for site specific mycorrhiza, or proof of mycorrhiza on the root structures. This causes me concern. Here is a list of problems.
1. Slides of the mycorrhizal roots are without meaning. If you do not know what species the mycorrhiza is, root slides are valueless. Moreover mycorrhizas can be soft as plants are. A mycorrhiza that is viable in a greenhouse or with high water/fertility is worth little or nothing in the wild. (It can i.d. out to the same species but not be hardy in the wild.)
2. Dictating to the grower how to grow mycorrhizal plants is laughable. We are tracking for each site specific species, (at present) 1200+ different variables/species. With the mainline species we only need to track only 5-10. These are proprietary and range from light intensity to mycorrhizal type.
3. We are hearing of many growers growing natives for the first time getting site specific projects. We are often called afterwards to re-grow the plant material to replant. You cannot grow natives as other plant material has been grown and have viable mycorrhiza.
Here is what I'd like to see required and why.
1. Require the grower to provide proof that he/she has grown at least plants(more than one) from that genera before. (On some weird things you may only find a grower that has grown that family before.)
2. Require the grower to provide a sworn statement that the stock has no fungicides or insecticides applied. Have an analysis done if it is questionable. Mycorrhiza do not tolerate either very well. (Other native growers, e.g., Native Sons, Yerba Buena, Theodore Payne, etc., do not use insecticides or fungicides past the propagation stage.)
3. Use a grower from an equal or harsher climate than the site. (Soft plants and mycorrhiza are unstable in a harsher site. The plants do not need to be site grown.)
"There are no silver bullets."(Trappe) Good ecology, botany, mycology and their use through horticulture are the primary tools we have.
Copyright 1995 Las Pilitas Nursery Santa Margarita - Escondido

39. Fungi and Douglas fir Forests 2 

Daniel continues to lead us into interesting and useful fields of thought. The value of fungi as wildlife food is underscored, as well as showing the critical value wildlife plays in maintaining a diverse fungal base, in turn prompting the trees to grow into the carbon machines the fungi need. The fungi secure minerals and store water via soil aggregation caused by glomalin deposited by the fungi.
I intend to continue posting my collected articles, especially concerning mycorhizzia, because it is a lot of dense but useful information relating to specifics of our original theme. From these two posts we get a good idea of the relationship. We will continue posting our knowledge base for fungi and Douglas fir, as well as explore the mechanics of the mycorhizzial partnership.
While I may appear to be going backward, from the landowners point of view this is the initial interest. Discovery of glomalin came years later for me enriching an already amazing tapestry of forest activity.
From: http://groups.google.com/groups?hl=en&lr=&ie=UTF-8&q=author:dwheeler%40teleport.com+ (dwheeler@teleport.com)
Subject: Mycorrhizal fungi importance
Newsgroups: bionet.agroforestry
Date: 1999/01/02
WARNING: This is not intended to be spam, but some may consider it so.
Last week I bought Chris Maser's Forest Primeval, an important work IMO for
all people working in either forest products or forest management. Although
the work is skewed to the Pacific Northwest, the interrelationships of
organisms will apply elsewhere too.
Maser notes in this 1989 book the importance of large (more than 3" diameter)
woody debris and its importance as animal habitat, truffle formation, water
reservoirs, and soil creation. Maser is a small mammal specialist, and was
responsible for getting the California Red-backed vole (Clethrionomys
californicus) OFF the Endangered Species List. In fact, Maser has called this
ubiquitous animal "the most common mammal west of the Cascades [Mountains]."
Several years ago Dr. James Trappe received a live vole late on a Friday. He
weighed the vole, placed it in a darkened cage (voles are so timid that
without cover, they literally die of fright) and placed an equal weight of
fresh truffles in with it, before locking up and going home. Early the next
morning he returned and found the vole dead. The truffles were gone. An
autopsy of the vole later showed it had died of starvation. This indicated a
vole must eat at least its body weight in truffles each day. While occasional
pieces of vegetation and probably some seeds are included in its diet, the
vole is the most abundant mycophagist in either forest or plantation, and is
one reason why trees continue to grow well in Oregon (at least in some
places).
Maser notes that truffles contain Clostridium butyricum, a "classic nitrogen-
fixing bacterium" and Azospirillum spp., a "nitrogen-fixing bacterium". When a
tree seed sprouts near truffle innoculum, the rootlets become quickly
innoculated with nitrogen-fixing bacteria as well as mycorrhizal truffles.
Maser goes on to note that Rhizopogon vinicolor, a common truffle frequently
collected near Douglas fir, is especially important to seedling tree survival.
Because of the increased absorption of water, phosphorus and potassium through
R. vinicolor mycelium, it greatly increases the health of its host tree, and
enhances the growth.
Last year I sold some R. vinicolor to create a slurry for a 50-acre
plantation containing slighly over 50,000 Douglas fir, Western hemlock, Grand
fir, and Ponderosa pine seedlings. As I recall, about 30-50 grams of dried
truffle were ground into paste, then a small amount was added to water
buckets into which the seedling roots were dipped. Last August I was invited
to visit the site to view the results.
It should be noted that in 1998 Oregon experienced an 83-day drought which
caused most fungi not to fruit.
Most of the trees were less than 20-inches tall, and had been planted in
stubble of an old wheat field. As I recall, most of the trees were planted in
late March and early April of 1998. In late August, 1998 I saw only two trees
which had not survived. I admit there were probably more. But the point is:
transplantation shock typically responsible for up to 15% mortality in
seedling trees locally, was minimized.
Using data supplied in Forest Primeval and The Redefined Forest (another book
by Maser), I calculated that a 20 gram vole consuming 20 grams of truffles
would contain approximately 9,000,000 spores. These spores would be
concentrated into 300 fecal pellets each day, containing about 300,000 spores
each. Since Maser considers this vole to be extremely abundant (between
500-3,000 individuals per acre), it is easy to see how new seedling trees get
inoculated with mycorrhizal fungi in nature.
What is less well known is that a single Douglas fir can be host to at least
50 mycorrhizal fungi at one time. The greater this mycorrhizal diversity is,
the more rapid trees are likely to grow. As individual trees mature, there
appears to be a change in the association of mycorrhizal fungi with the tree.
Few mycorrhizal fungi found with seedling trees are found in old-growth
forests.
In nature, according to Maser, truffles and other mycorrhizal fungi are
consumed by a plethora of animals including deer, elk, bear, voles,
squirrels, chipmunks, moles, gophers, mice, porcupines, pika, mountain
beaver, and other indigenous animals. At least 60 species of animals are
known to eat truffles on a regular basis based on analysis of fecal pellets.
(See Key to Spores of the Genera of Hypogeous Fungi of North Temperate
Forests with special reference to animal mycophagy, by Michael A. Castellano,
James M. Trappe, Zane Maser, & Chris Maser.)
In tree nurseries seedlings typically become innoculated with Hebeloma
crustuliniformis. However, according to work done in New Zealand, this fungus
is insufficient to grow trees rapidly. Until it is replaced, Douglas fir
seldom grows more than a few inches a year.
In 1986-89 I did several innoculation experiments at a Douglas fir Christmas
tree farm near Oregon City, Clackamas County, Oregon. One of the first
innoculations was with Rhizopogon vinicolor, R. parksii, R. villosullus, and
R. villescens, Suillus sps, Laccaria laccata, Laccaria
amethystina-occidentalis, Boletus zelleri, Boletus chrysenteron, and other
mycorrhizal fungi. Within 2 years of this multiple inoculation, most of the
trees were growing 3-8 feet per year! And where a 13-foot Douglas fir
Christmas tree was removed in Nov., 1990 and replaced by a 22-inch tall
4-year-old Douglas fir seedling (where Tuber gibbosum (Oregon White truffle)
was known to be fruiting) that seedling grew between February and October of
1991 at least 9.5 feet. That nearly allowed the tree to reach the height of
nearby trees. The following year it grew an additional 6 feet, and is now
(1998) nearly equal to its 40-60 foot tall neighbors.
Makes you kind of wonder how much tree cultivation is actually done, doesn't
it?
BTW, this stand was producing between 300-1300 pounds of Tuber (true
truffles) per acre in 1989-93, depending on the definition of species. Some
trees have several different species associated with them, allowing multiple
yearly cropping of fungi with the same trees, in addition to increasing the
tree growth rate exponentially. This suggests that fungi are worth far more
than the trees as timber. Each year. For 50 or more years.
I have some dried Rhizopogon vinicolor available, if anyone is planting
Douglas fir.

Daniel B. Wheeler
http://www.oregonwhitetruffles.com/

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38. Fungi in Douglas fir Forests I 

38. Fungi in Douglas fir Forests II have been searching the internet for several years looking for information relating to restoring our property. Daniel has demonstrated the critical importance, variety, abundance and economic potential of forest fungi in many postings. Daniel is convinced small landowners can profitably inoculate trees for annual returns valued in excess of the timber. There is so much here it is impossible to summarize, so I am going to paste a series of reports exactly as they appear on the internet for the education of the public, and to build the evidence that fungi produce tons of soil glue from carbon dioxide through production of glomalin, essential to water storage in the soil.
From: Daniel B. Wheeler (dwheeler@ipns.com)
Subject: Fungi as additional tree crops
Newsgroups: bionet.agroforestry, alt.forestry
Date: 2002-09-15 21:35:50 PST
In "Growing Gourmet and Medicinal Mushrooms" (c.1993, Ten Speed Press)
Paul Stamets states: "Most ecologists now recognize that a forest's
health is directly related to the presence, abundance and variety of
mycorrhizal associations."
Dr. James M. Trappe, professor emeritus at Oregon State University,
says "It is easy to find trees in the Pacific Northwest without
mycorrhizal fungi: look for trees without green."
Some 3,000 species of mycorrhizal fungi are known to associate with
Douglas-fir alone.
Perhaps 50 mycorrhizal fungi have ever been cultivated. Some of these
include Geopora cooperi, Lacterius deliciosus, Genea intermedia,
Gautieria monticola, Scleroderma cepa, S. aurantiacum, S. hypogaeum;
Pisolithus tintorius, Boletus chrysenteron, B. zelleri; Laccaria
laccata, L. amethystina-occidentalis; Leucangium carthusiana; Tuber
gibbosum var. gibbosum, T. g. var. autumnale, T. gilkeyae, T.
sphaerosporum, T. spinoreticulatum, T. sp. nov.; Helvella lacunosa,
Hymenogaster parksii, Hysterangium coriaceum, Truncocolumella citrina,
Barssia oregonensis; Rhizopogon parksii, R. vinicolor, R. villescens,
R. villosulus, and R. colossus. The author has "planted" the spores of
these all species, and found fruiting bodies close to where he
"planted" them. While not definitive, it may be cultivation.
Why are mycorrhizal fungi important to tree growers?
Mycorrhizal fungi gather water and soil nutrients, leach phosphorus
and potassium from rock, associate with nitrogen-fixing bacteria, and
may act as fungal prophylactics against soil pathogenic fungi.
Many mycorrhizal fungi are also expensive foods. These include the
commonplace Western chanterelle (Cantharellus formosus), the esteemed
Oregon Black truffle (Leucangium carthusiana), and the valuable
matsutake (Tricholoma magnivelare).
Paul Stamets (op cit) notes: "Mycorrhizal mushrooms form a mutually
dependent, beneficial relationship with the roots of most plants,
ranging from trees to grasses. ... both organisms benefit from this
association. Plant growth is accelerated."
These fungi also diversify income for tree growersand land owners. A
study in Southern Oregon with a stand of Lodgepole pine (Pinus
contorta) and matsutake (Tricholoma magnivelare) showed that the
mushroom production each year was more valuable than the trees as
timber.
My mycorrhizal fungi inoculation methods differs from those used to
grow the French Black truffle (Tuber melanosporum) by Truffe
Agronomique of France. Rather than inoculating seedlings in nurseries,
I inoculate acres of trees at a time. My method costs a lot:
$2500/acre (about 500 trees), but compared to buying inoculated
truffle trees, costs only one-third the cost. (See costs for T.
melanosporum trees at www.garlandgourmettruffles.com).
Ironically, Ian Hall in "The Black Truffle" (c. 1994, New Zealand
Institute for Crop & Food Research Limited) notes that most truffle
growers still view other mycorrhizal fungi as competitors to truffle
production. The opposite may be true.
Stamets (op cit) further states that "Fungi and their host trees may
have long associations without the appearance of edible fruitbodies."
In other words truffle mycorrhizae is not enough _by itself_ to
produce truffles.
The North American Truffling Society (PO Box 296, Corvallis, OR
97339) has accumulated several thousand collections of truffles. The
Oregon White truffle, Tuber gibbosum var. autumnale, is one of the
most common collections. But there are several other hypogeous fungi
which fruit at the same or different times of the year, and which are
common to Oregon White truffle producing lands. These species can be
found with younger trees. They suggest there is a successional
relationship among mycorrhizal fungi as trees age. Seedling trees (1-5
years old) have one or more of these five fungi. Oregon White truffles
are usually found among slightly older trees (5-50 year old trees).
Matsutake (Tricholoma magnivelare) were uncommon in the Mt. St.
Helens eruption zone until after 1995, but are becoming more common.
Chanterelles (Cantharellus formosus) are found with trees as young as
15 years old in the Coast range of Oregon, but are uncommonly found
with trees less than 400 years old near Timothy Lake in the Oregon
Cascades at 3500 feet elevation. King boletes (Boletus edulis) have
been found with young Lodgepole pine near Cape Lookout, Oregon. But
may associate with only 75 year old Sitka spruce in the same area
(personal observations, author).
While identification of mycorrhizae by root formation is still in its
infantcy, data published on the Internet has found 7 fungal species
colonizing _the same_ .5 cm rootlet! A typical seedling tree has
hundreds of sites. A healthy, 100-year-old tree may have several
million of those sites.
The demand for truffles and truffle products continues to rise in the
United States. Truffle producers including tree farmers may profit as
a result.
Because truffles are also mycorrhizal, trees associated with truffles
tend to grow more rapidly. A 22-inch tall seedling tree planted at
Paul Bishop's Jones' Creek Tree Farm grew to more than 10.5 feet
between February and October, 1991. Truffles were found with the
previous tree, and with this tree at a later date.
Helen V. Smith was an expert in Cortinarius mushroom identification,
all of which are believed to be mycorrhizal. She once found an
isolated Douglas-fir along the Oregon Coast in a sand dunes area that
was fruiting 53 species of Cortinarius _at the same time_.
Alexander H. Smith and Nancy Smith Weber, in "The Mushroom Hunter's
Field Guide" (c. 1958, 1963, 1980 by The University of Michigan)
states in the introduction that "A single tree may (on different
roots) form mycorrhizae with several species of fungi, and there is
increasing evidence that many of the plants in a forest are
inter-connected through these mycorrhizal networks in the soil. ...
one can predict where to find a species of mushroom by studying the
distribution of its host tree. Furthermore, since trees are generally
more conspicuous than mushrooms, knowing various tree species and
their associated fungi can help collectors locate likely habitats for
particular species."
It is possible to find many mycorrhizal fungi fruiting at different
times of the year with individual trees. Cultivation of these fungi
dramatically expands crops associated with trees. The author has found
Oregon White truffle (Tuber gibbosum var. autumnale), Oregon Gray
truffles (Tuber gibbosum var. gibbosum) and Oregon Black truffles
(Leucangium carthusiana) with the same trees at two different sites
within the same year. Sites with similar characteristics inoculated
with these truffles might reasonably produce multiple truffle crops.
How long do truffle-inoculated trees live? No one knows at this time.
Truffles have been found with trees as young as 6 inches tall and
perhaps 3 years old. Truffles have also been found with trees 8 feet
in diameter and probably 200 (or more) years old.
Glomalin is a protein exudate of some mycorrhizal fungi. It has been
found to sequester large quantities of CO2 for long time periods in
soils. (See http://www.ars.usda.gov/is/AR/archive/sep02/soil0902.htm)
This sticky protein causes soil particles to clump together, allowing
air and water to penetrate to greater depths and increasing soil
fertility.
Dr. James M. Trappe has noted that truffle mycelium (the plant-like
portion of truffles) also produces an exudate which causes soil
particles to clump together.

The above information cannot be complete. Too few fungi have _ever_
been examined for their gastronomic importance or their association
with trees, both in forests and in plantations. The subject in an
on-going topic of great interest to me.

Posted as a courtesy by
Daniel B. Wheeler
www.oregonwhitetruffles.com

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Thursday, July 01, 2004

Effects of Elevated CO2 on Tree Seeds  

36.Effects of Elevated CO2 on Tree Seeds
http://www.co2science.org/print.php3
Seeds (Trees) -- Summary

How does enriching the air with carbon dioxide impact the reproductive capacity of trees? LaDeau and Clark (2001) addressed this question in a major way when they determined the reproductive response of loblolly pine trees to atmospheric CO2 enrichment at Duke Forest in the Piedmont region of North Carolina, USA, where in August of 1996 three 30-m-diameter FACE rings began to enrich the air around the 13-year-old trees they encircled to 200 ppm above the atmosphere's normal background concentration, while three other FACE rings served as control plots. Because the trees were not mature at the start of the experiment they did not produce any cones until a few rare ones appeared in 1998. By the fall of 1999, however, the two scientists found that, compared to the trees growing in ambient air, the CO2-enriched trees were twice as likely to be reproductively mature and produced three times more cones per tree. Similarly, the trees growing in the CO2-enriched air produced 2.4 times more cones in the fall of 2000; and from August 1999 through July 2000, they collected three times as many seeds in the CO2-fertilized FACE rings as in the control rings.
Also working on this aspect of the Duke Forest FACE study were Hussain et al. (2001), who report that (1) seeds collected from the CO2-enriched trees were 91% heavier than those collected from the trees growing in ambient air, (2) the CO2-enriched seeds had a lipid content that was 265% greater than that of the seeds produced on the ambient-treatment trees, (3) the germination success for seeds developed under atmospheric CO2 enrichment was more than three times greater than that observed for control seeds developed at ambient CO2, regardless of germination CO2 concentration, (4) seeds from the CO2-enriched trees germinated approximately five days earlier than their ambiently-produced counterparts, again regardless of germination CO2 concentration, and (5) seedlings developing from seeds collected from CO2-enriched trees displayed significantly greater root lengths and needle numbers than seedlings developing from trees exposed to ambient air, also regardless of growth CO2 concentration.
What are the implications of these findings? The propensity for elevated levels of atmospheric CO2 to hasten the production of more plentiful seeds on the trees of this valuable timber species bodes well for naturally-regenerated loblolly pine stands of the southeastern United States, where LaDeau and Clark report these trees "are profoundly seed-limited for at least 25 years." Hence, as the air's CO2 content continues to climb, they conclude that "this period of seed limitation may be reduced," which is good news for this highly-prized tree. In addition, the observations of Hussain et al. suggest that loblolly pine trees in a CO2-enriched world of the future will likely display significant increases in their photosynthetic rates. Enhanced carbohydrate supplies resulting from this phenomenon will likely be used to increase seed weight and lipid content. Such seeds should consequently exhibit significant increases in germination success, and their enhanced lipid supplies will likely lead to greater root lengths and needle numbers in developing seedlings. Consequently, when CO2-enriched loblolly pine seedlings become photosynthetically-active, they will likely photosynthesize and produce biomass at greater rates than those exhibited by seedlings growing under current ambient CO2 concentrations.
Another major study of the reproductive responses of trees to elevated levels of atmospheric CO2 was conducted at the Kennedy Space Center, Florida, USA, where in 1996 three species of scrub-oak (Quercus myrtifolia, Q. chapmanii, and Q. geminata) were enclosed within sixteen open-top chambers, half of which were maintained at 379 ppm CO2 and half at 704 ppm. Five years later -- in August, September and October of 2001 -- Stiling et al. (2004) counted the numbers of acorns on randomly selected twigs of each species, while in November of that year they counted the numbers of fallen acorns of each species within equal-size quadrates of ground area, additionally evaluating mean acorn weight, acorn germination rate, and degree of acorn infestation by weevils. So what did they find?
Acorn germination rate and degree of predation by weevils were unaffected by elevated CO2, while acorn size was enhanced by a small amount: 3.6% for Q. myrtifolia, 7.0% for Q. chapmanii, and 7.7% for Q. geminata. Acorn number responses, on the other hand, were enormous, but for only two of the three species, as Q. geminata did not register any CO2-induced increase in reproductive output, in harmony with its unresponsive overall growth rate. For Q. myrtifolia, however, Stiling et al. report "there were four times as many acorns per 100 twigs in elevated CO2 as in ambient CO2 and for Q. chapmanii the increase was over threefold." On the ground, the enhancement was greater still, with the researchers reporting that "the number of Q. myrtifolia acorns per meter squared in elevated CO2 was over seven times greater than in ambient CO2 and for Q. chapmanii, the increase was nearly sixfold."
Stiling et al. say that these results lead them to believe "there will be large increases in seedling production in scrub-oak forests in an atmosphere of elevated CO2," noting that "this is important because many forest systems are 'recruitment-limited' (Ribbens et al., 1994; Hubbell et al., 1999)," which conclusion echoes that of LaDeau and Clark with respect to loblolly pines. If other trees behave similarly, it would appear that the rising CO2 content of earth's atmosphere will be a boon indeed to the regenerative prowess of the planet's forests. More research results from other tree species are thus anxiously awaited.
References
Hubbell, S.P., Foster, R.B., O'Brien, S.T., Harms, K.E., Condit, R., Wechsler, B., Wright, S.J. and Loo de Lao, S. 1999. Light-gap disturbances, recruitment limitation, and tree diversity in a neotropical forest. Science 283: 554-557.
Hussain, M., Kubiske, M.E. and Connor, K.F. 2001. Germination of CO2-enriched Pinus taeda L. seeds and subsequent seedling growth responses to CO2 enrichment. Functional Ecology 15: 344-350.
LaDeau, S.L. and Clark, J.S. 2001. Rising CO2 levels and the fecundity of forest trees. Science 292: 95-98.
Ribbens, E., Silander, J.A. and Pacala, S.W. 1994. Seedling recruitment in forests: calibrating models to predict patterns of tree seedling dispersion. Ecology 75: 1794-1806.
Stiling, P., Moon, D., Hymus, G. and Drake, B. 2004. Differential effects of elevated CO2 on acorn density, weight, germination, and predation among three oak species in a scrub-oak forest. Global Change Biology 10: 228-232.

Page printed from: http://www.co2science.org/subject/s/summaries/seedstrees.htm Copyright © 2004. Center for the Study of Carbon Dioxide and Global Change

Commentary: Here again we have evidence showing the advantages of heightened carbon dioxide levels for recovering forests. The trees are setting seeds earlier, the seeds are more viable and they have higher survival rates. CO2 Science Magazine does an excellent job attacking the science behind climate change arguments but it fails to recognize the critical issue of fungi storing carbon in the soil and its global impacts, or the consequences when this important relationship is ignored or neglected.
Opportunities to repeat these experiments with local trees present themselves. The concept of FACE experiments would seem like a good idea. One also wonders if enriching selected seed trees with CO2 for a boost to create more viable seedlings and give better results in planting out, especially in situations where the object is getting trees established, for instance, in erosion control or slide stabilization plantings.
This brings us back to our illusory Institute of Mountain Agriculture. What is the effect of elevated on grafted material? Will elevated CO2 provide better survival of hardwood grafts in forest trees? Will selected and grafted forest trees ever become a large scale reality? Will the growing power of hardwoods be harnessed by annual harvesting? What is the mycorhizzial relation in mixed forests? Plenty of questions, but slowly they are being refined.

36. Politics Delays Coho Protection Listing Again 

Governor's Appointee Is Blocked
State Senate leader acts after interim member of fish and game panel votes to delay protection for coho salmon. But she may yet retain her post.
By Kenneth R. Weiss, Times Staff Writer July 1, 2004
http://www.latimes.com/news/science/la-me-salmon1jul01,1,2336401.story?coll=la-news-science The battle over listing coho salmon took another turn yesterday as Governor Schwarzenagger’s choice for a post on the state Fish and Game Commission was blocked by Senate Majority leader Dan Burton after she voted to delay listing coho as an endangered species. Acting as an interim commissioner, Marilyn Hendrickson, co-owner of a fishing tackle outfit and vice president of a nonprofit group set up to work with state officials to enhance "angling opportunities in the state" and co-produces a "California Sportsmen" radio show, joined two other Davis appointees in voting to delay listing California coho to the state's list of threatened and endangered species. She was set to be confirmed Wednesday by the Senate Rules Committee, .when Dan Burton yanked her name from consideration. Burton said he was upset at her vote and would meet with her and straighten her out or he would find a new commissioner. Term limits mean Burton will leave the State Senate this year, while Hendrickson has until March to be approved. She said she is not worried in the least about it, and declined to give reasons for voting for the delay.
Last month Burton and Sher (Senate Environmental Quality Committee Chairman) sent all commissioners a letter urging they vote to list coho north of Punta Gorda threatened, and south of Punta Gorda as endangered. There are about 5,000 coho remaining from historical stocks of around 250,000. The letter said there was no scientific basis for delaying listing. Yet that is exactly what happened when commissioners met last week in Cvrescent City, ignoring recommendations by state Department of Fish and Game and National Marine Fisheries Service.
Commissioners Micheal Flores, James Kellogg and Hendrickson approved delaying the vote, continuing a pattern begun in 2000. Last week they postponed action on the commission's decision last February that directed state officials to begin the process of giving coho salmon protected status. Instead, they asked that state officials check whether federal officials might do more, given that the coho have some measure of protection under the federal Endangered Species Act. The delays are increasingly frustrating Burton and Sher. The two, together with the proponents of listing currently on the Board are convinced coho will disappear from California in the next ten years if protections are not put in place.
Opposing industries include timber, afraid of more restrictive forest practice rule; and cattle and alfalfa irrigators who resist withdrawal limits and oppose screens to protect juvenile fish. Farmers are afraid restrictions will take land out of agriculture and add expensive new rules, and possibly limit water use, according to Commissioner Flores, who said he was sympathetic, and that he hoped voluntary efforts would begin to make a difference. Nothing changes the fishes current status, illegal to catch commercially or by sportsmen since 1988.
Tom Weseloh, California Trout regional manager, said "The water in these streams is over-allocated so a lot of these streambeds go dry. Dry streams cannot grow fish." Commissioner Sam Schuchat and Commissioner Bob Hattoy lost last weeks vote. Schuchat said: “"I'm convinced that if we don't protect this species as endangered, it's going to go extinct in the next decade. It may already be too late."
Commentary: It is boggling to watch supposedly informed people deny old science repeatedly demonstrated to be true. If they can’t get over the old, how will they come to grips with the new? The timber industry has an opportunity now to think in terms of glomalin and sedimentation. Positive action by them in this area might really give coho a glimmer of hope for survival by improving habitat, or at least slowing its destruction. Glomalin gives a scientific basis and predictability for ground disturbance when it is ignored. Glomalin studies will eliminate clear cuts and should reduce road building but is the missing tool needed for sustainable land use practices.
Cattlemen and farmers need to start thinking about retaining precipitation on their own lands. Once the water is in a creek bed it is a public resource, and will always be subject of debate. We need farmers, and they need water, but not at the expense of entire industries, communities and species. Understanding glomalin allows farmers more flexibility in water management schemes and should reduce the need for withdrawals. It is a question of whether we are restoring a remenant of historical species or changing the landscape to one that can not support its current menagerie of biota- the beginning of a post-industrial ecology completely different from that which it sprang from.

Wednesday, June 30, 2004

35. UNEP talks shareholder values of Sustainability 

35. New UNEP Report Pinpoints Necessity of Environmental and Social Research to Protect Shareholder ValueThursday, June 24, 2004
UNEP
Twelve large global institutional investors today released a statement about the need for environmental and social issue awareness if large corporations are going to stay profitable in a changing world. Improvements in environmental and social areas lay the groundwork for successful businesses. Their statement backs Kofi Anan’s U.N. Global Compact, an independent evaluation of the corporate social responsibility initiative.
The summary report, titled "The Materiality of Social, Environmental and Corporate Governance Issues to Equity Pricing", warns long-term investment losses if the areas of environment and social issues are not addressed in an aggressive way. The twelve represent companies managing $1.6 trillion in assets.
Speaking from the Summit UNEP Executive Director Klaus Toepfer said: "This new report is a crucial recognition from major financial institutions that the environmental and social components of sustainable development, as well as the economic considerations, should sit at the heart of investment and capital market considerations."
"The financial analysts who undertook the research believe sustainability issues impact long-term shareholder value. It is clear, however, that to protect shareholder value the response must start with action today by companies serious about our environment and that wish to contribute to thriving communities worldwide"… .
The report was produced for the UN and covered eleven sectors in eight industries and was the first time the financial impact of environmental, social and corporate considerations and criteria were evaluated through the lens of industrial portfolio and pension plan outlooks.
Some industries like oil and gas, aviation, utilities and insurance already threatened with climate change, while others have developing opportunities in awakeneing carbon markets.
“Some of the key findings include:
* Environmental, social and corporate governance issues affect long-term shareholder value. In some cases those effects may be profound.
* Financial research is hindered both due to the paucity of reporting on the part of many companies concerning environmental, social and corporate governance issues and because of insufficient disclosure of these issues in annual reports.
* Financial research is greatly aided when there are clear government positions with respect to environmental, social and corporate governance issues. In some cases analysts were not able to provide in-depth reports due to a lack of certainty regarding government policy.
"The analyst findings demonstrate clearly that consideration of environmental, social and corporate governance factors are essential to prudent investment management and, therefore, essential to the fiduciary responsibility of pension fund trustees and investment managers", said Carlos Joly, Co-Chair of the UNEP FI Asset Management Working Group, and representative of Storebrand Investments. "It is to be expected that regulators will take this into account when updating fiduciary law and that institutional investment consultants will also take notice", he said.
The release coincides with a report by stock exchanges of their support for the principles of the UN Global Compact. Georg Kell, of the UN Global Compact Office said "financial markets are awakening to the fact that environmental and social issues have important financial impacts."

Copies of the report will be available mid-day on 24 June on the web at: www.unepfi.net/stocks
Information about the UN Global Compact can be found at:
www.unglobalcompact.org
For more information, contact:
Robert Bisset
Information Officer for Europe, UNEP
rbisset@unep.fr
Web site: http://www.unep.org/
Commentary: This is an important acknowledgement of the seemingly obvious. You can run neither your markets nor your resources into the ground and expect to continue to profit. Year to year analysts’ sheets tell nothing of depleted resources or restoration timetables. Underpaid labor does not create market bases. Walking away from depleted resource sites is a commonly accepted practice. Operations injurious to others need to be addressed.
This refocusing on a bottom line shadowed by social and ecological issues is a good sign for the future. Corporations will strive to protect their interests and gain advantages. Once the competition becomes a competition of profitable sustainability we will see some exciting changes.

Wednesday, June 23, 2004

Desertification 

33.Desertification
A conference in Bonn tomorrow will address desertification. As we can see in the article, this is a human caused condition created by people on the land. It is clear nothing is known about glomalin, that land preservation is not an option, nor are stricter laws concerning firewood cutting or burning for agriculture. It is clear no advantage is found in higher growth rates due to increased CO2, and that it is politically very difficult to leave land alone long enough to revegetate it to its former self.
As a result, we have JR Smith’s adage, “In men’s footsteps, the desert” occurring before our very eyes. Poverty and ignorance account for a good deal of the problem. Greed over water for cities and agriculture are depleting biologically necessary water. Vast landscapes have had their natural water regimes disrupted, leaving large areas dry and drying. Timber removal causes more runoff and less storage, shrinking of the soil moisture zone and surface drying. Local watercourses dry up in the heat. Lack of shade raises temperatures and evaporation rates. Precipitation storage shrinks and susceptibility to insects, disease and fire rise. Seedlings have a hard time surviving through summer. All the carbon in the vegetation and the soil is returned to the atmosphere as CO2, no more is returned. Soil loses its tilth and degrades back to its mineral components, easily blown or washed away, and filling streams and rivers with sediment.
Desertification archeology as a human caused phenomenon deserves close attention. An atlas of lost lands would be very interesting. One of Pacific Lumber s founders knew of a place in Maine where sand replaced soil after clear cutting so that no trees could grow back. He was afraid the same thing would happen around the Great Lake states when he was there. It helped form the philosophy that made the company sustainable through the years.
Increased temperature could exasperate places alreadt drying but it, together with elevated carbon dioxide are usable tools for remediation. We have these facts to encourage us:
Drip irrigation can allow us to establish and re-establish forests in drying lands with minimal water.
Elevated Carbon Dioxide reduces stoma cells, reducing evapo-transpiration
Shade reduces temperature, protects fungi. Mulch can be used around individual trees.
Shaded streams run all year
Glomalin production rises with higher heat and CO2
Glomalin binds small soil particles, trapping soil moisture and preventing erosion
Glomalin itself is the topsoil needed for sustainable productivity
Desertification means destruction of a regions glomalin zone

See Our Shrinking Watersheds, Redwood Reader #3, April 29, 2004
http://redwoodreader.blogspot.com/2004_04_25_redwoodreader_archive.html

World's land turning to desert at an alarming speed, warns United Nations
http://www.enn.com/news/2004-06-16/s_24932.asp
Wednesday, June 16, 2004
By Chris Hawley, Associated Press
UNITED NATIONS — The world is turning to dust, with land the size of Rhode Island becoming desert wasteland every year and the problem threatening to send millions of people fleeing to greener countries, the United Nations says.
One-third of the Earth's surface is at risk, driving people into cities and destroying agriculture in vast swaths of Africa. Thirty-one percent of Spain is threatened, while China has lost 36,000 square miles to desert — an area the size of Indiana — since the 1950s.
This week the United Nations marks the 10th anniversary of the Convention to Combat Desertification, a plan aimed at stopping the phenomenon. Despite the efforts, the trend seems to be picking up speed, doubling its pace since the 1970s.
"It's a creeping catastrophe," said Michel Smitall, a spokesman for the U.N. secretariat that oversees the 1994 accord. "Entire parts of the world might become uninhabitable."
Slash-and-burn agriculture, sloppy conservation, overtaxed water supplies, and soaring populations are mostly to blame. But global warming is taking its toll too.
The United Nations is holding a ceremony in Bonn, Germany, on Thursday to mark World Day to Combat Desertification and will hold a meeting in Brazil this month to take stock of the problem.
The warning comes as a controversial movie, The Day After Tomorrow is whipping up interest in climate change and as rivers and lakes dry up in the American West, giving Americans a taste of what's to come elsewhere.
The United Nations says:

* From the mid-1990s to 2000, 1,374 square miles have turned into deserts each year, an area about the size of Rhode Island. That's up from 840 square miles in the 1980s and 624 square miles during the 1970s.

* By 2025, two-thirds of arable land in Africa will disappear, along with one-third of Asia's and one-fifth of South America's.

* Some 135 million people — equivalent to the populations of France and Germany combined — are at risk of being displaced.

Most at risk are dry regions on the edges of deserts, places like sub-Saharan Africa or the Gobi Desert in China, where people are already struggling to eke out a living from the land.
As populations expand, those regions have become more stressed. Trees are cut for firewood, grasslands are overgrazed, fields are over-farmed and lose their nutrients, water becomes scarcer and dirtier.
Technology can make the problem worse. In parts of Australia, irrigation systems are pumping up salty water and slowly poisoning farms. In Saudi Arabia, herdsmen can use water trucks instead of taking their animals from oasis to oasis, but by staying in one place, the herds are getting bigger and eating all the grass.
In Spain, Portugal, Italy, and Greece, coastal resorts are swallowing up water that once moistened the wilderness. Many farmers in those countries still flood their fields instead of using more miserly drip irrigation, and the resulting shortages are slowly baking the life out of the land.

The result is a patchy "rash" of dead areas, rather than an easy-to-see expansion of existing deserts, scientists say. These areas have their good times and bad times as the weather changes. But in general, they are getting bigger and worse-off.
"It's not as dramatic as a flood or a big disaster like an earthquake," said Richard Thomas of the International Center for Agricultural Research in the Dry Areas in Aleppo, Syria. "There are some bright spots and hot spots. But overall, there is a trend toward increasing degradation."
The trend is speeding up, but it has been going on for centuries, scientists say. Fossilized pollen and seeds, along with ancient tools like grinding stones, show that much of the Middle East, the Mediterranean, and North Africa were once green. The Sahara itself was a savanna, and rock paintings show giraffes, elephants, and cows once lived there.
Global warming contributes to the problem, making many dry areas drier, scientists say. In the last century, average temperatures have risen over 1 degree Fahrenheit worldwide, according to the U.S. Global Change Research Program.
As for the American Southwest, it is too early to tell whether its six-year drought could turn to something more permanent. But scientists note that reservoir levels are dropping as cities like Phoenix and Las Vegas expand.
"In some respects you may have greener vegetation showing up in people's yards, but you may be using water that was destined for the natural environment," said Stuart Marsh of the University of Arizona's Office of Arid Lands Studies. "That might have an effect on the biodiversity surrounding that city."
The Global Change Research Program says global warming could eventually make the Southwest wetter, but it will also cause more extreme weather, meaning harsher droughts that could kill vegetation. Currently, the Southwest drought has become so severe that even the sagebrush is dying.
"The lack of water and the overuse of water: That is going to be a threat to the United States," Thomas said. "In other parts of the world, the problem is poverty that causes people to overuse the land. Most of these ecological systems have tipping points, and once you go past them, things go downhill."
Source: Associated Press

Tuesday, June 22, 2004

33. Elevated CO2: What Can It Do for Semi-Arid Shortgrass Steppe  

33. Elevated CO2: What Can It Do for Semi-Arid Shortgrass Steppe Vegetation?
Volume 7, Number 25: 23 June 2004

This appeared in this weeks CO2 Science magazine. They clearly show the root zone soil moisture improvements from elevated CO2 and warmer temperatures. CO2's influence on stomata is making plants more water efficient and resistant to ozone. Still no mention of glomalin although these scientists are from USDA SAS.
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How will earth's natural ecosystems respond to the ongoing rise in the air's CO2 content? This question is of paramount importance to the current debate over the nature of CO2-induced global change. Will plants and the animals that depend upon them for both sustenance and shelter be helped or hurt by the consequences of carbon dioxide's direct plant physiological impacts? And what of possible warming, CO2-induced or otherwise? Will it exacerbate deleterious effects or enhance good ones?
Nelson et al. (2004) broached these fundamental questions in a five-year study (1997-2001) of the semi-arid shortgrass steppe (SGS) of Colorado, USA. Working at the USDA-ARS Central Plains Experimental Range in the northern portion of the SGS about 60 km northeast of Fort Collins, Colorado, they used large (15.5 m2) open-top chambers to examine the effects of elevated CO2 (720 vs. 360 ppm) on plant water relations, ecosystem water use efficiency, soil moisture dynamics and root distributions of the ecosystem's dominant C3 (Pascopyrum smithii and Stipa comata) and C4 (Bouteloua gracilis) grasses. So what did they find?

The five Agricultural Research Service scientists and their collaborator from Colorado State University report that "seasonal average soil moisture throughout the soil profile (0-15, 15-45, 45-75, 75-105 cm) was increased under elevated CO2 compared to ambient CO2 for much of the study period," with the greatest relative increase (16.4%) occurring in the 75-105 cm depth increment. They remark that this finding of "increased soil moisture under elevated CO2 at the deepest soil depth suggests that water percolated deeper into the soil profile and that less moisture was lost to evapotranspiration under elevated CO2." Noting that "this phenomenon enhances water storage in the deep fine sandy loam soils underlying large portions of the SGS," they go on to say that "this increase in soil moisture has been shown to be the major controlling factor in improved carbon assimilation rates and increased total aboveground biomass in this system (LeCain et al., 2003) and will likely decrease the susceptibility of the SGS to drought."

Another important finding of the group of Colorado researchers was, in their words, that when averaged over the study period, "leaf water potential was enhanced 24-30% under elevated CO2 in the major warm- and cool-season grass species of the SGS (Bouteloua gracilis, C4, 28.5%; Pascopyrum smithii, C3, 24.7%; Stipa comata, C3, 30.4%)." They say these results are similar to those of "studies involving other C3 and C4 grass species (Owensby et al., 1993; Jackson et al., 1994)," and that the enhanced leaf water potential - "which reflects improved plant water status and increased drought tolerance (Tyree and Alexander, 1993)" - may lead to increased leaf turgor and allow the grasses "to continue growth further into periods of drought." Hence, it is not surprising that, averaged over the five years of the study, Nelson et al. found that "water-use efficiency (g aboveground biomass harvested / kg water consumed) was 43% higher in elevated than ambient CO2 plots."

In discussing the broader implications of their findings, the scientists say their results "suggest that a future, elevated CO2 environment may result not only in increased plant productivity due to improved water use efficiency, but also lead to increased water drainage and deep soil moisture storage in this semi-arid grassland ecosystem." And they say that "this, along with the ability of the major grass species to maintain a favorable water status under elevated CO2, should result in the SGS being less susceptible to prolonged periods of drought."

That Nelson et al.'s findings are the norm and not the exception is confirmed by their noting that "previous studies have reported increased soil moisture under elevated CO2 in semi-arid C3 annual grasslands in California (Fredeen et al., 1997), mesic C3/C4 perennial tallgrass prairie in Kansas (Owensby et al., 1993, 1999; Ham et al., 1995; Bremer et al., 1996), and mesic C3 perennial grasslands in Switzerland (Niklaus et al., 1998) and Sweden (Sindhoj et al., 2000)." Hence, we can validly expect the beneficent effects of atmospheric CO2 enrichment revealed in this impressive study to be found in grasslands throughout the world as the air's CO2 content continues to rise to double-and-beyond its current concentration.

But what if air temperature rises concurrently? Actually, things could get even better under that scenario. Nelson et al. note, for example, that "air temperature was on average 2.6°C higher inside the chambers than outside," and they say that this warming "was implicated in the 36% enhanced biomass production observed in chambered-ambient compared to non-chambered plots." Consequently, since this already-enhanced biomass production was the starting point from which the 41% increase in biomass elicited by the doubling of the air's CO2 content was calculated, the increase in biomass caused by the concurrent actions of both factors (increasing air temperature and CO2 concentration) could well be something on the order of 90%.

So bring on the climate alarmists' "twin evils" of elevated CO2 and temperature … and let the (ecological) good times roll.

Sherwood, Keith and Craig Idso

References
Bremer, D.J., Ham, J.M. and Owensby C.E. 1996. Effect of elevated atmospheric carbon dioxide and open-top chambers on transpiration in a tallgrass prairie. Journal of Environmental Quality 25: 691-701.

Freden, A.L., Randerson, J.T., Holbrook, N.M. and Field, C.B. 1997. Elevated atmospheric CO2 increases water availability in a water-limited grassland ecosystem. Journal of the American Water Resources Association 33: 1033-1039.

Ham, J.M., Owensby, C.E., Coyne, P.I. and Bremer, D.J. 1995. Fluxes of CO2 and water vapor from a prairie ecosystem exposed to ambient and elevated atmospheric CO2. Agricultural and Forest Meteorology 77: 73-93.

Jackson, R.B., Sala, O.E., Field, C.B. and Mooney, H.A. 1994. CO2 alters water use, carbon gain, and yield for the dominant species in a natural grassland. Oecologia 98: 257-262.

LeCain, D.R., Morgan, J.A., Mosier, A.R. and Nelson, J.A. 2003. Soil and plant water relations determine photosynthetic responses of C3 and C4 grasses in a semi-arid ecosystem under elevated CO2. Annals of Botany 92: 41-52.

Nelson, J.A., Morgan, J.A., LeCain, D.R., Mosier, A.R., Milchunas, D.G. and Parton, B.A. 2004. Elevated CO2 increases soil moisture and enhances plant water relations in a long-term field study in semi-arid shortgrass steppe of Colorado. Plant and Soil 259: 169-179.

Niklaus, P.A., Spinnler, D. and Korner, C. 1998. Soil moisture dynamics of calcareous grassland under elevated CO2. Oecologia 117: 201-208.

Owensby, C.E., Coyne, P.I., Ham, J.H., Auen, L.M. and Knapp, A.K. 1993. Biomass production in a tallgrass prairie ecosystem exposed to ambient and elevated CO2. Ecological Applications 3: 644-653.

Owensby, C.E., Ham, J.M., Knapp, A.K. and Auen, L.M. 1999. Biomass production and species composition change in a tallgrass prairie ecosystem after long-term exposure to elevated atmospheric CO2. Global Change Biology 5: 497-506.

Sindhoj, E., Hansson, A.C., Andren, O., Katterer, T., Marissink, M. and Pettersson, R. 2000. Root dynamics in a semi-natural grassland in relation to atmospheric carbon dioxide enrichment, soil water and shoot biomass. Plant and Soil 223: 253-263.

Tyree, M.T. and Alexander, J.D. 1993. Plant water relations and the effects of elevated CO2: A review and suggestions for future research. Vegetatio 104/105: 47-62.
a

32.Fire Goats and Opportunities 

32.Fire Goats and Opportunities
A request from a local fire-safe council for information about using goats for fuel load reduction led to a review of the literature. Several years ago I had started collecting information about this when I first saw it in connection with the Oakland Hills after the devastating fire there in 1991. Over the years several more articles appeared in print and on TV about using them for fire protection in the urban interfaces as well as traffic medians. The need to reduce fuel loads is reaching a critical point with vegetation responding to increased carbon dioxide and warmer temperatures with explosive growth.
An Agrarian History of Great Britain tells about the earliest settlers there, at about 7000 B.C., cleared the first lands for agriculture. The original forest had lots of elm, and elm leaves are pretty good forage, with a fair amount of nitrogen. The early method was to lop branches from the trunks and use the leaves as fodder. The trunks were left and became quite burled. These trees were cut by later inhabitants and used to form burial chambers in the long barrows. Many of these are lined with beautiful elm posts.
Archelogists reportedly have a hard time telling sheep from goats but sheep have to have grass. The history discusses the fact that research shows every single part of Britain had been covered with trees. Even the moors had been cleared in the past, but had not regrown as forest. Goats are as good a candidate as any for the first known deforestation, and goats have been used in woodlands since the dawn of time.
A search on google now brings about 340,000 hits in .13 seconds for fire goats. The Forest Service, Bureau of Land Management, Caltrans and many smaller public agencies are using goats in steep, difficult or brush covered terrains. There are all kinds of goat web sites with just a wealth of information on every aspect of raising, caring for and using goats.
Numbers always help us get a better picture of opportunities. Three hundred and fifty goats will clear an acre in a day. One price I saw was a dollar fifty per day per head, five hundred twenty-five dollars per acre per day with 350 goat herds.
The local need for fire goats is immense because the scale of fire prevention is vast, both wildland and at the edge of development. Enriched atmospheric CO2 and warmer temperatures will cause even more explosive vegetative growth. Increasing canopy height and removing fuel loads and fire ladders make a big difference in fire defense.
To the goat herd, this results in a continuous food source for goats. While some of the original programs I read about took any goat, specific breeds can be used for hair, hides, meat, some used for milk, some companies offer sheep for grazing certain types of grass. Opportunities abound for fire goats. Many jobs need doing, while the goats’ food source is expanding into increased fire danger. Even without contracts fodder is becoming more available as the products become better known.
Fire goats can be a tremendous help in restoring forest systems by controlling brush, Goats can prepare a site for planting if natural regeneration is insufficient. If it is, goats can be introduced after trees grow beyond browse height for fuel reduction and conifer release in some areas. Goats eat many noxious weeds and can be used to reduce problem plants like Scotch broom. Goats reduce roadside vegetation readily and can be used in residential interfaces, stewardship zones, firebreaks, and regenerating forestlands.
Goat Wisdom: Most everything you need to know about raising goats. http://www.goatwisdom.com

From California Grazing
http://www.californiagrazing.com/_fpclass/index.htm“Yellow star thistle is named for the bright, thistle like flower that have sharp spines surrounding their base. It is a long-lived annual and is found at elevations of 7000 feet or less. It grows to any where between 6 inches to 5 feet tall. Most of the plants seeds germinate within a year of disbursement, however some can stay viable for up to 3 years. Goat grazing is a highly effective way of reducing star thistle and star thistle seed production. Goats will eat the plants in all stages, including after the spines form. Surprisingly goats quite like thistle and when present it is one of over time by goats provides positive and successful results in the eradication of star thistle.
Aggressive noxious weeds like thistle bring problems as they displace beneficial plants, reduce habitat and recreational value. Goat grazing is also effective control of other weed species such as Spurge, Nettles, Purple Star thistle, Artichoke thistle, Poison Ivy and Poison Oak. Where as human contact with Poison Oak or Poison Ivy can cause a allergic reaction in humans, goats relish them and are highly effective at eradicating this weed. Goat grazing is a cost effective, ecologically sound way to clear land and promote growth of native grasses and beneficial plants.”

http://www.albrightseed.com/wildfires.htm
Wildfires Don’t Have A Goat of a Chance
© 2001 Wendy Dager
Despite their reputation, goats don't really eat tin cans.
But, oh, how they love weeds. And shrubs and forbs and grasses.
With proper control, goats and other animals with voracious appetites for greenery can be used to scale back the threat of wildfires, including those that could be rampant during the upcoming windy season in Southern California.
Often egged on by dry Santa Ana winds, 6,000 wildfires per year wreak havoc in California. Among the worst in history was the 1999 fire season when 273,000 acres and 300 homes were destroyed at a cost of $500 million. Such statistics are expected to worsen as the number of fires increases due to the rapid expansion of housing developments, which sprout ever closer to locations that are vulnerable to fire.
One of the hardest hit areas in the last decade was Oakland Hills, when a 1991 fire claimed more than 2,400 homes. Determined to keep it from happening again, the local government sought out alternatives to the few available preventive techniques, which include the more conventional herbicides and controlled burns.
Instead, Oakland officials called in a goat rancher, who provided the goateed, bleating, four-footed crew that happily chomped the fire-prone hillsides for two weeks at a hefty $15,000 per job.
"Theres some irony here", said S&S Seeds' Paul Albright. "Not too long ago, the government used to charge goatherders for grazing rights. Now, city governments are paying them to come in with livestock to clear the land."
According to Dr. An Peischel, those prior fees weren't fair at all to the person providing the goats, and, at times, the best management practices for grazing were not used.
"It's a bad precedent that people who live in cities and own land would charge farmers to graze their goats on them", said Peischel. "What happened years ago, is that farmers might abuse the land. They'd graze it improperly because they had to pay a lot of money for it."
Peischel knows what she's talking about. A PhD in Range Livestock Nutrition and a goat farmer for 18 years, she and partner Mike Spaetgens began their business, Goats Unlimited, in Hawaii. Their herd of goats was hired by growers of sugar cane, citrus, coffee, bananas, and papaya to clear land prior to planting, as well as to perform weed control duties between harvests.
Using the Kiko breed of goats, along with livestock guardian dogs to herd them, Peischel and Spaetgens main objective is to enhance land productivity.
"If you want your land well taken care of, then you better find a good rancher that's going to be a steward to your land", said Peischel. "As farmers, we're doing a landowner a service. We're preventing fires on their land and we're enhancing their perennial grasses so we're enhancing watershed management."
Now located in Rackerby, California, an hour-and-a-half north of Sacramento, Goats Unlimited is truly what it says it is: unlimited in the services it performs.
"We do all different kinds of things with the goats", said Peischel. "We do land rejuvenation, erosion control, restoration projects, fire breaks, and fuel load reduction. We provide breeding stock. We make meat sales to organic restaurants in San Francisco and Berkeley. We sell them to folks who have small farms - five, ten, fifteen acres - that want to do land cleaning so that their places don't burn on the urban/wildlife interface. We clear ditches for irrigation companies so that the water flows freely and you dont have a lot of weeds and stuff along the banks."
Currently, the Goats Unlimited herd numbers 700, but each spring it expands to between 1200 and 1300 head. The care and feeding of the goats includes supplementing their diet with something other than that which they cull from the land.
"If you're doing a fire break in an old ponderosa pine forest, there's not much to eat there", said Peischel. "If nutrition is lacking, protein has to be supplemented. Though utilizing livestock to manage land isn't new, goats have been an industry in California for only five years. Their use, however, is becoming more widespread as fire prevention and mitigation practices evolve."
"There are various tools to mitigate or minimize the damage done by fire to grasslands, rangelands, forests, homes and personal property", said Peischel. "Each tool has a specific use and place in management."
Weed abatement tools include the mechanized variety such as bulldozers, masticators and chipping equipment.
Using machines, however, is sometimes hazardous they can spark and cause fire. Which is why, in June of 2001, the city of Sunnyvale, California employed goats to maintain local landfills. According to an article by Gretchen Knaup of the Sunnyvale Sun newspaper, one of the reasons the goats were used was because the many pipes and wells in the landfills were difficult for tractors to get around and there was risk of starting a vehicle fire.
Peischel admits that an employer has to be receptive to the idea of fire control via hooved herbivores.
"We have to find people that want to pay us to do this", she said. "Proper planning, site evaluation and the working of the goats takes time. Contract price depends on the size of the job; if your'e doing fire breaks; how old the goats are; what's the weather; what's the vegetation."
"Each contract", says Peischel, "is individually negotiated, and consists of coordinating a variety of sources, including the local fire patrol, professional fire abatement teams, California Department of Forestry, and others." Regardless of the number of parties involved and the combination of factors that are unique to each job, Peischel emphasizes that the purpose remains how to best utilize the goats to decrease the amount of fuel that may cause a wildfire.
"The aim is to break the continuity of flammable cover, creating defensible space", said Peischel. "Once an area has been brushed by the goats, it can be maintained as a living green belt."
Peischel is pragmatic about her unusual business, but believes the goats are here to stay and that providing them for land management is the career for her.
"If I had to go to work, I don't know what I'd do", she joked. "I just can't imagine having a real job." For more information, visit http://www.goatsunlimited.com or call (530) 679-1430.

Living Systems Land Management uses goats for fire mitigation, habitat restoration, erosion control and vegetation management.
Visit http://www.lslm.com.

Monday, June 21, 2004

31. Hybrid Owl Species Complicates Bird Future  

31.Hybrid Owl Species Complicates Bird Future
http://www.newsday.com/news/science/wire/sns-ap-spotted-owl-hybrid,0,3507314,print.story?coll=sns-ap-science-headlines
By JEFF BARNARD Associated Press Writer
June 21, 2004, 2:09 AM EDT
LOWELL, Ore. -- It hoots kind of like a northern spotted owl, and looks kind of like a northern spotted owl. And like a spotted owl, it swoops in to take a mouse offered on a stick by U.S. Forest Service scientist Eric Forsman in a rainy stand of old-growth Douglas fir on the Willamette National Forest. However, this is a hybrid -- a cross between a northern spotted owl and a barred owl -- and it is one of the wrinkles in the future of the bird that triggered sharp logging cutbacks in the Northwest in 1994.
The invasion of the barred owl into spotted owl territory over the past 30 years and creation of the hybrids has become the top issue in the review of Endangered Species Act protection for the northern spotted owl, granted in 1990 largely due to loss of its old growth forest habitat to logging.
A panel of experts will report Tuesday in Portland on new information gathered for the U.S. Fish and Wildlife Service, which must make a decision by Nov. 15 on whether to maintain threatened species listing for the spotted owl. The latest studies show spotted owls are still declining, though just why remains a big question. Loss of old growth forest habitat has been minimal, particularly on federal lands where logging is restricted. Meanwhile, the barred owl is pushing spotted owls out of the way when it moves in.
"Clearly the barred owl is having more of an impact on the spotted owl than any of us anticipated 10 years ago," said Jerry Franklin, a University of Washington forest ecology professor serving on the panel. "The question now has to do with how much that impact is going to be. Is the barred owl essentially going to drive the northern spotted owl out of part of its range?"
The timber industry, which called for the review, argues that if barred owls push spotted owls out of old growth forests, those stands no longer have to be left standing as habitat, unless someone is willing to start killing barred owls. "It seems like the original basis for listing is really in question at this point," said Ross Mickey, western Oregon manager for the American Forest Resource Council.
Conservationists counter that protecting old growth forests may be more important than ever with the invasion of the barred owl. “The barred owl was around at the time of the listing," said Susan Ash, conservation director for the Audubon Society of Portland. "It's reached the radar screen to the point that, yes, it's a new threat. The numbers are high. "But nobody has an explanation for why they have come into the area. And nobody can prove they are actually causing an impact to owl numbers in the long term. This may be some natural process where two species figure out their own roles in the ecosystem."
Barred owls began moving west from forests in eastern Canada and Minnesota in the early 1900s. After reaching southwestern British Columbia, they moved south, appearing in spotted owl territory in Washington in 1973 and Oregon in 1978, according to a paper by Forsman and Oregon State University graduate student Elizabeth Kelly. They now reach into Northern California.
Barred owls are bigger and more aggressive than spotted owls, and there is evidence they sometimes kill their smaller cousins. Barred owls nest in the same kinds of places -- cavities in large trees -- and eat the same kinds of things, small rodents like flying squirrels and woodrats.
There is no good overall population estimate on barred owls or spotted owls, but when the two come together, the smaller and meeker spotted owl generally loses, though not always, Forsman said. "In a lot of study areas in Oregon, even though we are seeing gradually increasing numbers of barred owls, the spotted owl population seems to be holding relatively stable or only declining slightly," Forsman said. "So it's still up in the air what this is going to mean long term."
Cross breeding remains rare -- only 47 hybrids have been confirmed in the wild, mostly in Oregon -- probably due to behavioral differences between the two. "It probably occurs in most cases in a situation where there's a dearth of potential mates for the barred owl," Forsman said. "But that we don't understand very well."
On the Net: Owl links: http://www.nps.gov/olym/hand/owllinks.htm.

Commentary: It can always be expected for people to want to cash in on resources and that the ability to generate cash flow by extracting resources will always imperil existing conditions. Not being fully cognizant of all aspects of an old growth forest would seem to be enough reason to protect old growth that remains. The northern spotted owl has given us a reprieve until new information should come available. That information, in the form of understanding mycorhizzial produced glomalin, soil based carbon storage and its impact on soil moisture, soil stability, and stored carbon dioxide are among the vital reasons to protect old growth in and of itself. Habitat must be preserved and created for threatened, endangered and extirpated species to return to the surrounding landscape.
It is less clear what, if anything we can discern about the owls struggle(?) between species. Adaptation is one of the basic tools of natural selection. Territorial competition and expansion as conditions favor one species over another is being played out all over the world in the form of invasive and exotic species. Many find niches with no natural controls and proliferate to the point of harm, often crowding out natives. Changing temperatures in many environments mean they are recovering in conditions different from what they originally adapted to. Time may help sort it out but the rule seems to be if a new species is well adapted with advantages over natives, they will win the day. And too often we don’t see the danger until the new species is too well established to handle easily.


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