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  2002 Table of Contents

New Insights into Management of Soilborne Crop Pathogens
Under Direct Seeding: Take-all

R. James Cook, Endowed Chair in Wheat Research, Departments of Plant Pathology and
Crop and Soil Sciences, Washington State University, Pullman, WA 99164-6430

Take-all, caused by the soilborne pathogen Gaeumannomyces graminis var. tritici, is a root and crown disease of wheat, barley, and related forage and weedy grasses. The disease was first observed in the Pacific Northwest in 1901 at Albany, Oregon-one century ago!. The pathogen is thought to be native to this region. Until the last 25 to 30 years, take-all was known primarily if not exclusively on wheat under irrigation in southern Idaho, Yakima Valley, and Columbia Basin, and in high-rainfall areas such as the Willamette Valley of Oregon and western Washington. In contrast, the disease was observed in the dryland Pacific Northwest (PNW) in earlier years only in the wettest parts of the region, such as the Camas Prairie and eastern Palouse, and then only in years of exceptionally high rainfall. However, we have continued to see evidence of a wider distribution of take-all than has been assumed, and this was recently confirmed.

A survey conducted by graduate student Nathan Ramsey during 1998, 1999, and 2000 revealed that, of 270 PNW wheat fields planted to winter or spring wheat, where wheat had been grown either the previous year (high risk) or 2 years earlier (medium risk), 75% had enough take-all to be limiting to yield in the year it was sampled (Ramsey, 2001). There was no difference in the amount of take-all whether the field had been cropped to wheat the previous year or if the last wheat crop was 2 years earlier. Since most rotations in the Inland Northwest include winter or spring wheat every other year, 2 years in 3, or every year, we now recognize take-all as one of the most important diseases limiting yields of wheat in the dryland PNW.

"Dryland" versus "wetland" take-all

Take-all develops quite differently on wheat under dryland conditions compared to wheat grown in high-precipitation or irrigated areas. Since the classic, "textbook" description of take-all is based on how the disease develops under wet conditions, the disease has been over looked or "under recognized" in the dryland PNW-until recently.

Under wet conditions, such as in western Oregon and western Washington, and under irrigation, the disease develops as patches of stunted and prematurely blighted plants (white heads), with the most severe stunting in the centers of the patches where the plants were infected first. With suitably moist soil, the pathogen grows from root to root and plant to plant radially outward from centers of infection in the field. Plants infected last, i.e., those plants on the edges of the patches, show the least or not stunting, although they may still ripen slightly prematurely to produce shriveled grain. The other classic symptom of "wetland" take-all is the black mycelial growth of the pathogen on and within the seminal and crown roots, into the crown, and 1-2 inches up the stem. This diagnostic growth up the stems, which some call the "black stocking," can be seen by pealing back the leaf sheaths to reveal lower-most internodes of the elongated stems.

Under dryland conditions, which includes the great majority of the nonirrigated PNW most years, the disease occurs on individual plants rather than plants in patches, and the growth of the fungus from roots upward usually stops in the crown where the tiller bases are connected to the mainstem. There is no black stocking symptom with dryland take-all. The "window" of suitably moist soil when the soil temperature is also favorable (40-60o F) is too narrow to permit more than limited growth of the pathogen from plant to plant or upward on the stems. Under these conditions, every plant must be infected separately from primary inoculum in the soil. On the other hand, the fact that crown infection is limited to the site where stem bases of tillers are attached to each other and to the mainstem is still enough damage to restrict flow of water up the stem and hence can still cause premature blighting (white heads). However, under dryland conditions, with each plant infected separately from primary inoculum, we find blighted plants next to healthy plants much as happens with Fusarium crown rot, Cephalosporium stripe, and Pseudocercosporella foot rot.

Wetland take-all is easy to recognize by the patches of plants with white heads and the black stocking at the lower-most internodes. Dryland take-all can be recognized by cutting the crown open with a sharp knife to expose the bases of the stem at the point where they are joined together and to the mainstem. With Fusarium crown rot, this tissue is typically brown, but with take-all, it is grey or grey with black mottling.

Management of Take-all with Crop Rotation

Historically, a 1-year break out of wheat or barley to a broad leaf crop or to oats or corn (both non host grass crops), has been considered sufficient for management of take-all. Again, this has been based on experience with "wetland" take-all where, once the crop is harvested, the moist conditions of the top soil while also still relatively warm favors breakdown of infected roots and tiller bases used by the pathogen as a food base to infect the next crop. Under dryland conditions in the PNW, the infected roots and tiller bases last for long time in the soil, and may last even longer when kept near the soil surface as with mulch-tillage and no-tillage. In other words, the drier the area in our region, the lower the risk of take-all in general but also the more likely that the pathogen inoculum will carry over in a dormant condition through a 1- or 2-year break.

Years ago, keen observers in Australia noted that take-all was most severe in years of a dry fallow prior to planting and a wet growing season after planting. The dry fallow delays or prevents microbial breakdown of the infested roots and crown tissues so that the inoculum potential of the pathogen remains high in spite of no hosts, while the wet growing season favors maximal infection and secondary spread from root to root, plant to plant, and up the stem.

The ability of the take-all pathogen to survive through 2 years of broadleaf crops in a low precipitation zone of the PNW is revealed by results from a direct-seed cropping systems study on the Ron Jirava farm west of Ritzville and led by William Schillinger. This study compared three cropping systems: continuous spring wheat; a 2-year spring wheat/spring barley rotation, and a 4-year safflower/yellow mustard/spring wheat/spring wheat rotation. In the 4th year of this study, the amount of take-all, although not severe, was the same on spring wheat whether continuous spring wheat, spring wheat after spring barley, or 1st or 2nd year spring wheat after back-to-back broad leaf crops (Table 1; Cook et al, submitted). Neither safflower nor yellow mustard is hosts of the take-all pathogen, and grass weed hosts were fully controlled. These results can only be explained on the basis of dormant carryover of the pathogen in infested wheat residue though 2 years without a host.

Table 1

Based on these observations, we can expect that a 1- or 2-year break to a nonhost crop will be useful in take-all management mainly or exclusively in areas with annual precipitation higher than 12 inches, but we are not sure how much higher. Tom Swaintz asked me to look at one of his wheat fields that had take-all, near Reardon in a 16-inch precipitation area. Two years earlier, one part of this field had been cropped to spring barley and the other part had been cropped to a broadleaf crop before the entire field was planted to two consecutive crops of wheat. Take-all was present throughout the field in this 2nd year of wheat, but was distinctly more severe, to the line, where spring barley had been grown 2 years earlier compared with the part of the field cropped to a broadleaf crop 2 years earlier. Under these conditions, the benefits of the broadleaf crop in the rotation extended into a second wheat crop. The chances of this outcome presumably would also depend on variations in precipitation from year to year and therefore year-to-year differences in buildup of the pathogen on host crops and the opportunity for dormant survival of the pathogen in infested roots and stem tissues between host crops.

With convincing evidence that the take-all pathogen can survive 2 years without a host in low-precipitation areas, the most reliable data on value of alternative crops in a rotation with wheat for take-all management are from the higher precipitation areas. Ramsey (2000, 2001) tested the benefits of spring barley compared with a broadleaf crop rotated with wheat in 2-year rotations in his survey of 270 fields conducted over the years 1999, 2000 and 2001. Focusing on fields in areas with > 18 inches of precipitation annually, the average take-all index for wheat after spring barley was 18.9 compared with an average of 11.2 for wheat after a broad leaf crop. This difference was significant at P = 0.5. The take-all index for wheat after wheat in this same precipitation zone was 16.0, i.e., essentially the same as for wheat after spring barley.

Independent studies indicated that a take-all index of 5 was the threshold index for take-all severity, above which take-all begins to have an affect on the yield. Thus, while spring barley was like another crop of wheat in these higher precipitation areas, take-all was still evident on wheat in 2-year broadleaf crop/wheat rotations.

It is worth noting that, on barley, the take-all pathogen tends to remain restricted to roots and extends only modestly or not at all into the crown or up the stems of adult plants, even under conditions favorable to "wet land" take-all. Because root tissues are short lived compared with cereal stem bases as a food base for the take-all pathogen, a 1-year-break between barley and wheat (i.e., spring barley/broadleaf crop/wheat) will likely provide better take all control than a 1 year break between wheat crops (i.e., wheat/broadleaf crop/wheat). Along with control of Cephalosporium stripe, this fact can help explain the success of the classic 3-year spring barley/pea or lentil/winter wheat rotation in the Palouse. This rotation provides 2 full years between wheat crops, for decomposition of wheat stem bases infested with the take-all pathogen, and 1 full year between barley and wheat, for decomposition of barley roots infested with the take-all pathogen.

Another advantage of spring barley over spring wheat prior to winter wheat is the slightly longer time (1-2 weeks or longer) between harvest and planting, owing to the faster maturity of barley. The longer the time between harvest and planting, the more time for death and disappearance of the take-all pathogen through decomposition of its food base, provided that soil conditions are suitably warm and moist for decomposition of cereal residue. The 7-8 months between harvest and planting a spring cereal is even better and can help explain some of the success in the higher rainfall areas of continuous spring barley or spring barley/spring wheat systems. Making the most of this system depends on early elimination of volunteer and grass hosts of the take-all fungus (the green bridge), much as has been shown for management of Rhizoctonia root rot.

Management of Take-all With Continuous Cereals

Rotation with a broadleaf crop, where ever possible, should always be the first choice for management of take-all in the higher rainfall areas. However, where broadleaf crops are not economical, take-all can still be managed while growing continuous cereals. This is because populations of rhizosphere bacteria suppressive to this disease build up in response to outbreaks of take-all and continuous wheat or wheat/barley sequences. The result is that take-all increases with continuous wheat or wheat/barley during the initial years, peaks out, and then declines in severity while yields recover. This cycle is known the world over as "take-all decline."

Take-all decline has only been documented in situations where take-all can or has become severe, i.e., in areas with wetland take-all or at the wetter extremes of areas with dryland take-all. No studies have been done to determine whether the soil also becomes microbiologically suppressive to take-all in the low precipitation zones of the PNW. Because the microorganisms responsible for take-all decline are, themselves, dependent on soil water potentials at or near field capacity (Howie et al.1987), it is doubtful that these bacteria represent a potential biological control of dryland take-all and even more doubtful that they can survive at effective populations during a year of fallow. The normal pattern observed in the dryland PNW is for wetland take-all to appear uniformly over the field in the second and sometimes the first year that the field is irrigated and annually cropped to wheat after decades of wheat fallow. This early and rapid development of take-all in response to irrigation is further evidence that the pathogen is already widely distributed in the field. Take-all decline then follows.

I have documented the rise and decline of take-all in two plots planted continuously over many years to wheat or wheat and barley. One was conducted at on the WSU station at Lind with winter wheat grown under irrigation and conventional tillage after decades of conventional management in a winter wheat/fallow sequence. The other was conducted on the Palouse Conservation Field Station near Pullman with alternating direct seeded spring and winter wheat. In both cases, take-all decline was complete by about the 15 year of monoculture. There were obvious improvements in yield already by the 10th to 12th year, but yields were disastrously low during the years of most severe take-all. In the experiment at the WSU stations at Lind, for example, yields in the 7th year of continuous wheat were only half of the yields in fumigated plots used as checks, and half of the yields achieved without fumigation in the 15th year of continuous wheat.

The natural drop in soil pH in response to ammonium forms of nitrogen is thought to favor or accelerate take-all decline. In this regard, we must pay particular attention to the possibility of more take-all if or when soils are limed.

One final point on management of take-all of continuous cereals: assuring adequate phosphorus availability for the crop can greatly help to limit the amount of this disease. P-deficient wheat is intrinsically more susceptible to take-all. With the loss of roots caused by the disease, the ability of the plants to obtain phosphorous is also then limited, making the plants more susceptible and aiding the pathogen to become even more aggressive as a parasite. Phosphorus placement below the seed or with the seed at planting, as done with direct seeding, can help over come this situation

Basic Research on Take-all Decline

Most of the basic research aimed at explaining take-all decline has been done at WSU by ARS scientists David Weller and Linda Thomashow with their students and postdoctoral associates. This work represents some of the most elegant research ever conducted in soil microbiology. Briefly, they have shown that continuous wheat and the consequent development of take-all favors a shift in make-up of rhizosphere bacteria towards a select group highly adapted to the rhizosphere of wheat and genetically equipped to produce an antibiotic, 2,4-diacetylphloroglucinol (PHL), inhibitory to the wheat take all fungus (Raaijmakers et al., 1997). The ability to make this antibiotic is a highly conserved trait in soil- and rhizosphere-inhabiting bacteria world wide, but different strains with this ability are adapted to the roots of different crops. The strains adapted to roots of wheat represent a recognizable genetic type that can now be quantified with a DNA-based test. Using this test, Raaijmakers and Weller (1998) showed that a population of about 105 of these bacteria per gram fresh weight of root is sufficient for suppression of take-all. The addition of only about 103 bacteria per seed, is enough to establish 105 per gram of root in the rhizosphere. This shows the remarkable ability of these bacteria to multiply in the rhizosphere of wheat. Most soil bacteria require an initial inoculum dose of 107 or 108 on the seed in order to achieve a population of 105 in the rhizosphere.

Field trials conducted with these rhizosphere-competent strains have produced modest but significant yield increases (Cook et al., submitted; Mathre et al.1999). Moreover, these strains are fully compatible with the chemical seed treatments currently used in the PNW (see Pythium report in these proceedings). Even better, these strains add value to the current chemical seed treatments, producing 8-10% greater yields and up to 15% greater yields than non-treated seed.

A combination of early elimination of the greenbridge, placement of fertilizer and especially phosphorous under the seed at planting, and treatment of the seed with a combination of chemical and biological seed treatments, has been shown to elevate yields in direct-seeded cereal-monoculture fields to within 90% of the yields obtained in these same fields in fumigated plots. Without the biological component, the yields are 80-85% of those in fumigated plots. Thus far, however, no company has been willing to take on this biological control technology for treating seed.

Is Take-all Favored by Direct Seeding?

A great deal of research has been done to determine whether or not take-all is worse with direct seeding, but the results have been mixed. In Georgia, where take-all is a problem in the wheat/soybean double crop system, the disease was more severe with conventional tillage and seeding compared with direct seeding (Rothrock, 1987). In contrast, research by graduate student Kevin Moore carried out with irrigated wheat in the 1970s on the WSU station at Lind revealed in side-by-side plots that take-all was more severe with direct seeding that when planting into a prepared seed bed (Moore and Cook, 1984). Similarly in Australia, side-by-side plots showed more server take-all with direct seeding than in plots tilled prior to seeding (Roget et al, 1996). It follows that infected plant remains left undisturbed in the soil will present a higher risk for infection of the next crop than when this tissue is fragmented into smaller pieces with tillage-like making kindling out of fire wood. It also follows that direct seeding is like moving the field into a higher precipitation zone and this we know will favor take-all. On the other hand, tilling the soil will also distribute the infested crop residue more uniformly so that more roots of the next crop will be exposed more uniformly to infection.

Ramsey (2001) looked at tillage effects on take-all in his 3-year survey of 270 wheat fields planted to wheat either the previous year or 2 years earlier. He found no difference in the severity of take-all between conventional and direct seeding. For example, of fields cropped the previous year to wheat, the average take-all index was 13.5 for conventional and 13.2 for direct-seeded fields. Focusing on high-precipitation areas (>18 inches annually), the average take-all index was 19.4 for conventional systems and 14.1 for direct-seed systems. The take-all indices for wheat from fields in areas with <18 inches annually were 10.6 for conventional and 11.9 for direct seeding. None of these differences were significant statistically.

Ramsey's results did not account for likely differences in fertilizer placement between conventional and direct-seed systems. Virtually all direct seeding includes phosphorus placement below or with the seed at planting, whereas few conventionally managed crops are fertilized at the time of planting. Possibly this difference in use and placement of phosphorous helps offset any additional inoculum potential of the take-all pathogen in direct seeded fields.

Paired-row Spacing

Widening the rows opens the wheat canopy to greater warming a drying of the top few inches of soil-conditions less favorable to take-all. However, widening the rows also reduces the plant population needed for maximum yield. The Yielder drill provides a way through a 5/15 inch paired-row spacing to have it both ways: open the canopy on one side of each row while providing the same plant density achieved with a uniform 10-inch row spacing..

Research conducted over several years with a plot drill that permits the openers to be either paired in a 7/17 inch spacing or spaced uniformly on 12-inch spacing indicates that, indeed, the amount of take-all can be limited significantly by pairing the rows (Cook et al, 2000). These studies included experiments with a fertilizer shank in each seed row so as to remove fertilizer placement as a variable. All studies were conducted in fields cropped to wheat 4 or more previous years and therefore were considered high risk for take-all. In addition, the studies were conducted in high-precipitation or irrigated areas so as to further increase the risk of take-all.

Two of the five studies that compared paired-row spacing with uniform spacing are described here as illustrations. One was conducted in an irrigated field near Pendleton and the other was conducted in a subirrigated site (bottom land) near Colfax. Although the experiments were conducted in different years in the mid 1990s, both sites were in the 9th year of continuous wheat when the respective experiments were conducted. Also, both sites were burned just prior to laying out and planting the plots.

At the Colfax site, fumigation was included as a treatment, and straw was returned to individual plots, so as to provide all combinations of burned + fumigated with no straw returned, burned + fumigated with straw returned onto burned-fumigated plots, burned but not fumigated, and burned, not fumigated, with straw returned. Each of these treatments was then seeded as side-by-side drill strips of paired and uniform row spacing with four replications. Yields averaged 95 to 100 bu/A with both paired and uniform row spacing in all fumigated plots, with our without straw returned to the plots, and in the nonfumigated treatment left bare-black (no straw returned to the soil). In contrast, returning straw to the plots after burning resulted in an average yield of 72 bu/A with uniform row spacing and 82 bu/A with paired-row spacing. Take-all mainly, but some Rhizoctonia and Pythium root rot, were severe in nonfumigated plots when covered with straw, and under these conditions, pairing the rows resulted in 10 bu/A additional yield of winter wheat.

We have consistently seen a 20-30 bu/A yield increase with winter wheat in response to burning in the high-precipitation areas of eastern Washington, but have also shown consistently that this yield response is nullified by returning straw to the burned soil surface immediately after planting (Cook and Haglund, 1991). In one such experiment near Pullman, we showed that the average percentage of adult winter wheat plants with seminal roots destroyed by take-all was 32, 18, 40, and 12, respectively, in replicated plots not burned, burned, burned and straw returned, and not burned but fumigated. These results collectively present a clear picture of less take-all and more yield in response to burning, but simply returning straw to the soil surface nullifies the burn effect. In other words, to the extent that the burn response is due to less root disease, the effect is probably through the warming and drying effect of making the soil surface bare-black and is not due to killing the pathogen in soil by the burn.

The experiment at Pendleton included paired- and uniformly spaced rows of winter wheat over alternating 24-foot-wide strips of tilled (disked twice) and not tilled land with stubble burned. Wheat planted into the tilled plots had less take-all and averaged 9 bu/A more compared to wheat in the direct seeded plots (109 versus 100 bu/A in tilled and no-till treatments respectively). These results support the work that suggests that take-all is favored by direct seeding, where every other management factor is held constant across the two treatments. When all data for tilled and no-till plots were combined, pairing the rows had no effect on yield but resulted in a 25% reduction in white heads and an increase in grain test weight of 0.5 lb/bu, both significant at P = 0.05. The effect of pairing the rows was real but subtle, being expressed as increased test weight but not increased yield.

One disadvantage of opening the canopy through paired-row spacing is great potential for weeds to grow in the space not planted to wheat. However, with new technology for weed control, especially with the introduction of herbicide-resistant varieties, paired-row spacing deserves a close look as one more tool in management of take-all and other root diseases without burning the straw.

Take-all Prediction and Risk Assessment-a Look at the Future

With knowledge of such variables as the recent (past 3-4 years) cropping history, time in weeks or months between harvest and planting, time between spraying and seeding for greenbridge control, and annual rainfall, it will be possible to develop a take-all prediction and risk assessment program for the PNW. Such a program is currently under evaluation in South Australia (David Roget and Michael Krause, personal communication) in combination with a DNA-based test to also provide the information on amount of inoculum of the take-all pathogen in the soil (Ophel Keller and McKay, 2001). The DNA-based test is proprietary with the South Australian Research and Development Institute (SARDI) and licensed to C-Qentec Diagnostics, a subsidiary of Aventis CropScience. Farmers can obtain a report on the amount of inoculum of G. graminis var. tritici, Rhizoctonia solani AG 8, cereal cyst nematode, two species of lesion nematodes, and both Fusarium pseudograminearum and F. culmorum, for about $A4 per acre. The results for fungal pathogens are expressed as pico grams (pg) of DNA of the target pathogen per gram of soil. For the nematodes, the results are expressed as numbers of nematodes (or eggs) per gram of soil.

In March, 2001, soil samples from 124 GPS-referenced sites representing 30 acres on the WSU Cunningham Agronomy Farm were sent to SARDI in Adelaide, Australia, for analysis of the complete package of pathogens for which their tests apply. The test was positive for the take-all fungus in only 16 of the 124 samples assayed, and these were low at 25 pg or less of DNA per gram of soil. Fifteen pg DNA/g soil is below the level of detection. These low numbers were surprising, considering that the field had been in spring barley in 2000 and spring wheat for several consecutive years before that. We must now test whether the field may be into take-all decline as occurred in the long-term direct-seeded plot on the nearby Palouse Conservation Field Station. It is also possible that the long period of no plants between harvest of the spring barley in 2000 followed by planting spring wheat in 2001, achieved with excellent greenbridge control in the fall of 2000, provided time needed for natural disappearance of the inoculum of this pathogen from the soil. Root assessments confirmed that take-all occurred in only trace amounts on the spring wheat in 2001, and the yield of 64 bu/A was respectable, considering the lack of moisture during the 2001 growing season.

We will be sending more soil samples from sites intended for direct seeded crops in 2002 in an effort to gain more experience with this technology and possibly help guide its introduction into use by growers in the PNW.

Conclusions

In spite of a century of research on take-all, this disease continues almost unabated, being subject as much or more to variations in the natural environment and the natural enemies than to anything we do by way of crop management aimed at its control. We now recognize that take-all management by crop rotation is achievable in the higher- but apparently not the lower-precipitation areas where the pathogen survives in crop residue easily through a year of fallow and even through 2 years of non host crops. Even in the higher-precipitation zones of the dryland PNW, including areas with 22-24 inches of precipitation annually, a 1-year break to a broadleaf crop helps but will not be long enough to lower take-all below an economic threshold. One possible exception is the 3-year rotation of winter wheat/spring barley/broadleaf crop, which allows the necessary 2 years between wheat crops. Barley, being a host of take-all, is like another crop of wheat when preceding wheat. However, because the pathogen tends to remain confined to roots of barley, and not extend into the stems, and because roots decompose faster than stems, the 1-year break to a broadleaf crop after spring barley and before returning to winter wheat can provide a significant rotational benefit for take-all control.

The results on whether or not take-all is favored by direct seeding have been mixed. When take-all is studied in side-by-side plots of till and no-till, take-all has consistently be more severe with no-till (direct-seeding) in both the PNW and Australia. This is thought to result from a higher inoculum potential of the pathogen in soil left undisturbed, at least within the old stubble rows, and also from the wetter and cooler soil environment in fields left covered with crop residue.

On the other hand, tilling the soil mixes the inoculum of the pathogen more uniformly into the soil. Further, direct-seeding is more likely than conventional seeding to include phosphorous placed directly under the seed or with the seed when planting, and availability of phosphorous is well known to limit the amount of take-all. Side-by-side comparisons allow for testing of a single variable such as till no not till, but direct seeding is a system that will likely involve a different planting date, different timing and method of greenbridge control, different method and timing of fertilization, different depth of seeding, and different row spacing, possibly paired-row rather than uniform-row spacing. These differences combined with low soil disturbance will determine the extent to which take-all will flourish or remain in check. Ramsey's survey shows that take-all is surprisingly common in conventional systems and that, taking all factors into account, both systems, because they include frequent crops of wheat, obviously are favorable to take-all.

Growers in the low-precipitation area can take solace in the fact that, while rotations and take-all decline are of little or no use for take-all management in their fields, take-all itself is limited by the low precipitation and will not be more than a mild or chronic disease problem, even with continuous spring cereals. Growers in the higher-precipitation areas can count on benefits from breaks to broadleaf crops as one approach, or from take-all decline as another approach. Since broadleaf crops tend also to disrupt take-all decline, consideration should be given to the use of either crop rotations that include broadleaf crops, or continuous cereals, but to not alternate between the two systems. Basic research on take-all decline continues with the goal of achieving biological control more consistently when applying the bacteria to seeds and thereby attracting a commercial interest willing to develop these bacteria into a product.

Literature Cited

Cook, R. J. and Haglund, W. A. 1991. Wheat yield depression associated with conservation tillage caused by root pathogens in the soil not phytotoxins from the straw. Soil Biol. & Biochem. 23:1125-1132.

Cook, R. J., B. H. Ownley, H. Zhang, and D. Vakoch. 2000. Influence of paired-row spacing and fertilizer placement on yield and root diseases of direct-seeded wheat. Crop Science. 40:1079-1087.

Cook, R. J., Schillinger, W. F. and Christensen, N.W. submitted.. Rhizoctonia root rot and wheat take-all in diverse direct seed spring cropping systems. Plant Disease

Howie, W. J., R. J. Cook, and D. M. Weller. 1987. The effect of soil matric potential and cell motility on wheat root colonization by fluorescent pseudomonads suppressive to take-all. Phytopathology 77:286-292.

Mathre, D. E., Cook, R. J., and Callan, N. W. 1999. From discovery to use-Traversing the world of commercializing biocontrol agents for plant disease control. Plant Disease 83:972-983.

Moore, K. J. and R. J. Cook. 1984. Increased take-all of wheat with direct-drilling in the Pacific Northwest. Phytopathology 74:1044-1049.

Ophel-Keller, K. and McKay, A. 2001. Root disease testing service: Delivery and commercialization. Page 17-18 in I. J. Porter, ed. Proceedings, Second Australian Soilborne Disease Symposium, March 5-8, Lorne, Victoria, Australia.

Raaijmakers, J. M., Weller, D. M. and Thomashow, L. S. 1997. Frequency of antibiotic producing Pseudomonas spp. in natural environments. Appl. Environ. Microbiol. 63:881-887.

Raaijmakers, J. M. and Weller, D. M. 1998. Natural plant protection by 2,4-diacetylphloroglucinol-producing Pseudomonas spp. in take-all decline soils. Mol. Plant-Microbe Interact. 11:144-152.

Ramsey, N E. 2001. Occurrence of take-all on wheat in Pacific Northwest cropping systems. M.S. Thesis. Washington State University, Pullman, WA.

Roget, D. K., Neate, S. M. and Rovira, A. D. 1996. Effect of sowing point design and tillage practice on the incidence of Rhizoctonia root rot, take-all and cereal cyst nematode in wheat and barley. Aust. J. Exp. Agric. 36:683-93.

Rothrock, C. S. 1987. Take-all of wheat as affected by tillage and wheat-soybean double cropping. Soil Bio. Biochem. 19:307-311.

     
 

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