British Columbia Mine Reclamation Symposium

Establishment and growth of mycorrhizal and rhizobium inoculated high-elevation native legumes on an.. Smyth, Clint R. 1997-07-02

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Proceedings of the 21st Annual British Columbia Mine Reclamation Symposium  in Cranbrook, BC, 1997. The Technical and Research Committee on Reclamation ESTABLISHMENT AND GROWTH OF MYCORRHIZAL AND RHIZOBIUM INOCULATED HIGH-ELEVATION NATIVE LEGUMES ON AN UNAMENDED COAL MINE SPOBL DUMP IN SOUTHEASTERN BRITISH COLUMBIA. C.R. Smyth, P.O. Box 517, Blairmore, Alberta, T0K 0E0. Tel: (403) 562-7083, Fax: (403) 562-7084, email: niyosotis@canuck.com. ABSTRACT Direct seeding, transplant survivorship, growth and reproductive performance of seven mycorrhizal and Rhizobium  inoculated high-elevation native legume species were studied at a coal mine in the Rocky Mountains of southeastern British Columbia. Mycorrhizal and Rhizobium  infection levels were low in the control treatment and highest in the combined inoculation treatments of all species. Transplant survival of each species was greatest for the combined inoculation treatment and lowest for the uninoculated control. Inoculation with mycorrhizae, Rhizobium or a combination of mycorrhizae and Rhizobium resulted in greater mean plant diameters, mean plant heights, mean numbers of leaves per plant and mean percentage flowering than the uninoculated control plants. Measured plant parameters for the single inoculation treatments, i.e., mycorrhizal or Rhizobium, were greater than the control, but the greatest increases were recorded for the combination inoculation treatment. The greatest differences in the measured parameters were recorded for silky locoweed (Oxytropis sericea Nutt.) although differences were also large for Bourgeau's milk-vetch (Astragalus bourgovii A. Gray), Robin's milk-vetch (Astragalus robbinsii [Oakes] A. Gray) and bent-flowered milk-vetch (Astragalus vexilliflexus var. nubilus Barneby). The results of the study have important management implications for the successful establishment of these species on unamended spoil dumps at high elevations. INTRODUCTION Agronomic legume establishment is problematic for high-elevation disturbances (Errington 1979, Johnson and Rumbaugh 1986, Smreciu 1993). Legumes or other species possessing a di-nitrogen fixation symbiosis have important ecosystem functions which can increase the nitrogen capital of the soil (Brown and Chambers 1990, Kenny and Cuany 1990) and stimulate biological soil activity (Flueler and Hasler 1990). Furthermore, legumes are important because of high levels of crude protein, vitamins and minerals (phosphorous and calcium) which make it valuable for forage and for di-nitrogen fixation (Klebesadel 1971). Grasses grown with legumes may benefit also from the nitrogen ?fixed? by legumes through re-cycling of decomposition and comminution of legume plant material (Chapin 1983). Success or failure of legume establishment on mine spoil is dependent upon several biotic and abiotic factors, not the least of which, is the legume - Rhizobium symbiotic system (Skousen 1986). The effective nodulation by appropriate Rhizobium species is essential (Somasegaran and Hoben 1994). Since mine spoil as a growing medium for plants is very infertile, three important factors must be considered: (1) what is the rhizosphere colonizing ability of a particular strain of Rhizobium, (2) how effective are the resultant nodules and (3) how does inoculation compare with the host plants' response to soil nitrogen. As well, mycorrhizal symbioses are known to enhance seedling establishment and survival (Allen and Allen 1980, Lambert and Cole 1980, Allen 1984, Allen et al. 1987, Waaland and Allen 1987) and plant growth (Hayman 1986, Azc?n-Aquilar and Barea 1992) through increased uptake of phosphorous and improved water relations or drought resistance (Allen and Friese 1992). The primary benefit of mycorrhizae is improved uptake of slowly diffusing ions of macronutrients such as phosphorous and 32 Proceedings of the 21st Annual British Columbia Mine Reclamation Symposium  in Cranbrook, BC, 1997. The Technical and Research Committee on Reclamation micronutrients such as zinc and copper (St. John 1990). Mycorrhizal symbionts are known to improve significantly inorganic nutrient uptake and growth in plants of boreal and alpine habitats (Mullen and Schmidt 1993). Mycorrhizal infection is also generally best developed under conditions of nutrient stress (Smith and Read 1996). In legumes, the efficiency of phosphorous nutrition appears to stimulate nodule production and increase the rate of atmospheric nitrogen fixation (Barea and Azc?n-Aquilar 1983, Barea et al. 1989). Authors studying Hedysarum boreale Nutt., a similar legume species to those of the present study, have indicated dependence of that species upon the mycorrhizal relationship for growth and survival (Reeves et al. 1979, Redente and Reeves 1981). Many of the species of this study have similar attributes and the potential for mycorrhizal infection is high. Therefore, inoculation of known mycorrhizal species may be a pre-requisite for establishment of these species on the low nutrient and moisture status spoil materials of waste rock dumps. RESEARCH OBJECTIVES The objectives of the study were to assess the efficacy of Rhizobium and mycorrhizal inoculation of a select group of native high-elevation legumes under growth chamber, glasshouse and field (mine spoil) conditions. THE STUDY AREA Based on the biogeoclimatic classification system used in British Columbia (Pojar et al. 1987), the study area for the Line Creek was located within the Engelmann Spruce - Subalpine Fir (ESSF) zone. The climate is characterized by short, cool growing seasons and long cold winters. The field experiments were positioned on a 5? west-facing slope of a dump platform (1930 m A.S.L.) dump platform. All research trials were established on unamended spoil. Composite (n = 5) spoil samples were withdrawn from each site, and their physical and chemical (nutritional) properties  analyzed  (Table 1). The spoils were composed primarily of siltstone and sandstone. In general, the spoils were high in percent coarse fragment contents (>2 mm) and percent sand (grain size <2 mm) and low in total nitrogen and available phosphorous.  At each location, spoil surface and interstitial resistances were checked with a hand held penetrometer and were found to be acceptable for seeding or transplanting. 33Proceedings of the 21st Annual British Columbia Mine Reclamation Symposium  in Cranbrook, BC, 1997. The Technical and Research Committee on Reclamation MATERIALS AND METHODS Species Selection Seven high elevation native legumes were selected for experimentation based on the research of Bell and Smyth (1988): alpine milkvetch (Astragalus alpinus L.), Bourgeau's milk-vetch (Astragalus bourgovii A. Gray), Robin's milk-vetch (Astragalus robbinsii [Oakes] A. Gray), bent-flowered milk-vetch (Astragalus vexilliflexus  var.  nubilus  Barneby), stalked-pod crazyweed (Oxytropis podocarpa A. Gray.) and silky locoweed (Oxytropis sericea Nutt.). Seed Collection and Sampling Seeds of each legume species were hand-collected from subalpine and/or alpine plant populations (metapopulations) located on selected mountains adjacent to each transplant location. All seeds were stored in paper bags at room temperature until extraction and cleaning. Once processed, the seeds were stored in plastic bags in a refrigerator at 2-5?C (Young and Young 1986).Seeds from a single accession of each species were withdrawn randomly with a Boerner Separator (Copeland and McDonald 1985). Single accessions were chosen to reduce within-species transplant survival variability. Accession selection was restricted to populations for which a large quantity of seed had been collected (N>1000). Seeds of each species were x-rayed for two minutes in a Faxitron 43855a X-Ray Unit set at 15 KVP (Smyth 1987). Pure Live Se S) were removed following examination of the x-rays. The seeds were surface sterilized in 4% Ca(OCl)2 for 10 minutes, rinsed five times in sterile water, soaked in 95% ethanol for 5 minutes and then rinsed a further five times (Garvin and Lindemann 1983). Seeds which appeared undamaged after the sterilization process were then scarified (single cut with sterilized scalpel). Rhizobium Inoculation - Greenhouse A four treatment randomized-block design experiment with three replicates (n=100) was performed to examine the efficacy of indigenous strain Rhizobium  inoculation on container seedlings. The four treatments were as follows: (1) control - uninoculated seedlings and fertilizer minus nitrogen, (2) uninoculated seedlings and fertilizer plus nitrogen, (3) Rhizobium  inoculation and fertilizer minus nitrogen and (4) Rhizobium inoculation and fertilizer plus nitrogen. A randomized block design (Little and Hills 1978) was employed. Seedling production involved several stages. Cone-Tainers? (depth - 16.3 cm, volume - 65.5 cm3) were surface sterilized with 2.5% NaOCl and rinsed three times in sterile water. Sterilized (autoclaved) growth medium of 2:1:1 (peat, vermiculite and perlite) was prepared and placed within the container cells. Two seeds per cone were placed on top of the soil mix and covered with one centimeter of sterilized forestry grit. The soil mix was then saturated with sterile Hoagland's solution (Hoagland and Broyer 1936) with or without nitrogen. The plants were inoculated with appropriate Rhizobium strains (liquid culture) after the cotyledons had cleared the surface of the vermiculite. Inoculation was delayed until after the emergence of the seedlings in order to eliminate the possibility of contamination. The cones were also covered with aluminum foil to prevent contamination during the first days of germination. Subsequent waterings alternated between sterile water and sterile Hoagland's solution (Garvin and Lindemann 1983). The plants were grown in a greenhouse until harvest after 4 months of growth (Gibson 1980). In the harvesting process, the Cone-Tainers? were inverted and the root system washed in a bucket of water to remove the growth media. The roots were then examined carefully. The respective Rhizobium cultures were grown in shake flasks of yeast extract-mannitol (YEM) broth for two weeks. An approximate initial population of 100 million rhizobia per inoculum bag was obtained in 34 Proceedings of the 21st Annual British Columbia Mine Reclamation Symposium  in Cranbrook, BC, 1997. The Technical and Research Committee on Reclamation order to ensure inoculation success. Rhizobium inoculum populations were estimated with the Most Probable Number (MPN) method of Vincent (1970). Equal volumes of liquid inoculum and sterile peat were mixed aseptically in whirl-pak bags and incubated at room temperature for 7 days, following which, the inoculum was stored in a refrigerator at 5?C until inoculation. The sources for the initial inoculum culture were fresh nodules from each species obtained during the previous summer. The data recorded was as follows: number nodulated, nodule location and plant growth. The distribution codes of Allen and Allen (1981) were used to classify species nodulation (Table 2).  Rhizobium Inoculation - Mine Spoil Commercial peat was purchased from the Nitragin Incorporated (Milwaukee, Wisconsin), divided into seven portions and sterilized in separate autoclavable bags. The peat in each bag was then re-inoculated with an indigenous strain of Rhizobium specific to each legume species. Ten fifty seed replicates which contained surface sterilized (refer to preceding section) seeds of each species were then coated with their respective strain of peat inoculum. Five replicates of each species were also coated with sterile non-inoculated peat. The replicates were then sown in a completely randomized block design at Line Creek during early spring. Seed was sown with an aluminum hand seeder which was sterilized between replicates with 70% ethanol. The developing seedling received weekly liquid fertilizer applications of Hoagland's solution (Hoagland and Broyer 1936). Five randomly selected replicates of inoculated seed of each species received a nitrogen Hoagland's solution treatment. Seedlings were monitored for growth and vigor. Seedlings growth and vigor were assessed visually and after two growing seasons, 100 plants per species and treatment were excavated to determine nodulation and infectiveness. The data recorded was as follows: number nodulated, nodule location and plant growth. The distribution codes of Allen and Allen (1981) were used to classify species nodulation (Table 2). Mycorrhizal Relationships Root samples were collected from three populations within the designated study area of the Southeast Kootenay Coal Block. Root samples were obtained by excavating five plants, carefully removing the adhering soil from the roots and excising 10 root segments from each plant. The root segments, 10-20 mm in length were then washed with distilled water, cleared stained and mounted in glycerine jelly (Phillips and Hayman 1970). All samples were then examined with a compound microscope to indicate presence or absence of fungal inf ection. No attempt was made to identify the fungus association or 35 Proceedings of the 21st Annual British Columbia Mine Reclamation Symposium  in Cranbrook, BC, 1997. The Technical and Research Committee on Reclamation quantify the amount of root infection. The fungal associations were classified according to the definitions presented in Moore-Landecker (1982). Mycorrhizal Inoculation A four factor experiment was performed to examine the effects of fungal associations on the survival, growth and reproductive activity of the legume species. The various factors were as follows: (1) control -1:2 ratio (volume) sterilized quartz sand and sterilized soil, (2) plus Rhizobium  treatment - 1:2 ratio sterilized quartz sand and sterilized soil plus Rhizobium inoculum, (3) plus fungi treatment - 1:2 ratio sterilized quartz sand and soil, and (4) + fungi and Rhizobium treatment -1:2 ratio sterilized quartz sand and soil. The soil for the plus fungi treatment was obtained by sterilizing an appropriate field soil for each species and then re-inoculating with spores and hyphae generated from greenhouse host "trap" plants (Schenck 1982). Two hundred plants per species and treatment were grown in sterilized Cone-Tainers? in the greenhouse for four months. The seedlings were then randomly assigned and transplanted into rows within blocks on a leveled area of the 1930 m dump. The seedlings were spaced twenty-five centimeters apart (Brockwell et al. 1982) within each block. Each block was separated by a one meter pathway. Dormant seedlings were transplanted by shovel shortly after snow-melt in the spring. Only healthy and nodulated seedlings were transplanted. Appropriate indigenous Rhizobium inocula were used for each species. All transplants were checked for viable Rhizobium nodulation prior to transplantation. As well, twenty-five plants of each species per treatment were randomly selected for destructive examination of fungal infection assessment prior to transplanting (Hayman et al. 1981). Plant height (cm), plant diameter, number of leaves, plant survival and flowering data was recorded after two years of growth. One hundred plants per species and treatment were randomly selected and excavated for assessment of mycorrhizal and Rhizobium  infection at the termination of the field experiment. The roots were examined following the procedures of Phillips and Hayman (1970) and classified according to the criteria of Carpenter and Allen (1988). No attempt was made to identify the fungus association or quantify the amount of root infection. RESULTS Rhizobium Inoculation - Greenhouse The growth chamber results are listed in Tables 3 and 4. The number of nodules per ranged from a mean of 9.2 for Hedysarum sulphurescens to 28.5 for Astragalus robbinsii. The percentage of effective nodules ranged from 57.8 for Astragalus vexilliflexus var. nubilus to 75.2 for Hedysarum sulphurescens. The distribution and number of effective nodules for each species was classified. Most of the nodules on each species were few in number and located on the secondary peripheral roots. Astragalus alpinus was an exception with several nodules located on both the crown as well as the peripheral root system.  36Proceedings of the 21st Annual British Columbia Mine Reclamation Symposium  in Cranbrook, BC, 1997. The Technical and Research Committee on Reclamation  Rhizobium Inoculation - Mine Spoil The mine spoil results are also listed in Tables 3 and 4. The number of nodules per plant ranged from a mean of 8.2 for Oxytropis podocarpa to 31.4 for Astragalus robbinsii. The percentage of effective nodules for each species ranged from 71.2 for Astragalus bourgovii to 95.4 for Astragalus vexilliflexus var. nubilus. Most of the nodules on each species were few in number and located on the secondary peripheral roots. In general, the number of effective nodules was greater on the mine spoil in comparison to the greenhouse. Modulation was sporadic for the uninoculated control. Astragalus alpinus and Astragalus robbinsii were not nodulated, whereas nodulation and nodule effectiveness was variable for the remaining species. Modulation and nodule effectiveness was high. Plants of Astragalus alpinus, Astragalus vexilliflexus var. nubilus and Oxytropis sericea were all inoculated and possessed effective nodules. The remaining species were inoculated, but the nodules were frequently ineffective. All plants of the inoculated plus nitrogen treatment were inoculated, but the percentage of effective nodules per plant was variable.Species growth and vigor was good or excellent for the inoculated and inoculated plus nitrogen treatments, but poor or fair for the uninoculated treatments (Table 5).Mycorrhizal Relationships All the selected species demonstrated some form of association with fungi. However, only Hedysarum sulphurescens was infected with vesicular arbuscular mycorrhizae. Mycorrhizal spore numbers among plant roots of Hedysarum sulphurescens was generally low although levels of root infection were high. External hyphae and often spores of the mycorrhizal fungi were common on the surfaces of all roots; however, mycorrhizal infections were only common on the smaller lateral roots. All stages of infection were observed: hyphal coils, simple hyphae, arbuscules and vesicles. Arbuscules were most frequent in the innermost cortical cells. Occasionally, primary and secondary lateral roots contained small amounts of minimally branched hyphae and scattered vesicles. VAM infection levels were qualitatively different between sample populations of Hedysarum sulphurescens, with infection levels greatest in the alpine populations. Evidence of dematiaceous septate fungi (DSF) was sporadic for the alpine populations of these species and non existent for the lower elevation populations. 37 Proceedings of the 21st Annual British Columbia Mine Reclamation Symposium  in Cranbrook, BC, 1997. The Technical and Research Committee on Reclamation  The Astragalus and Oxytropis species were only infected with dematiaceous septate fungi (DSF). The septate fungi were observed on the surface of the roots of these species and did not appear to have penetrated the root cortex. The walls of the hyphae were deeply melanized and septate. A number of clamp connections were observed as well. Clamp connections were more frequent with Astragalus vexilliflexus var. nubilus and Oxytropis sericea. All populations of the Astragalus and Oxytropis species were infected with (DSF), although there was an apparent increase in infection in populations found in higher elevation and exposed habitats. One hundred percent of the samples of collected for each of the Astragalus and Oxytropis species were associated with dematiaceous septate fungi. Mycorrhizal Inoculation - Mine Spoil Fungal and Rhizobium infection levels recorded during the final assessments were low for the control treatment for all species. The greatest fungal infections (Class 3) were recorded for the dual inoculation and fungal treatments although some of the transplants in the control and Rhizobium treatments had small amounts of fungal infections (Class 1 or 2) (Table 6). 38 Proceedings of the 21st Annual British Columbia Mine Reclamation Symposium  in Cranbrook, BC, 1997. The Technical and Research Committee on Reclamation Transplant survival of each species was greatest for the dual inoculation treatment and lowest for the uninoculated control. However, the differences in survival between treatments were not significant. One hundred percent of the dual inoculation treatment transplants of each species survived whereas, generally, one to two of the transplants of the single inoculation treatments died. All of the Astragalus alpinus transplants for each treatment survived.  Inoculation with mycorrhizae, Rhizobium or a combination of mycorrhizae and Rhizobium resulted in greater mean plant diameters, mean plant heights and mean numbers of leaves per plant than the uninoculated control plants (Table T). Measured plant parameters for the single inoculation treatments were greater than the control, but the greatest increases were recorded for the dual inoculation treatment. The greatest differences in the measured parameters were recorded for Oxytropis sericea although differences for Astragalus bourgovii, Astragalus robbinsii and Astragalus vexilliflexus var. nubilus were also large. Total above ground biomass and leaflet drymass biomass measurements were not recorded because bighorn sheep (Ovis canadensis) had grazed on the plants following the recording of the preceding measurements. Reproductive activity was also greatest for the dual inoculation although not significantly (Table 8). The control and the single inoculation treatments were very similar for all species except the Oxytropis species where there was a slight increase in the fungal inoculation treatment. 39 Proceedings of the 21st Annual British Columbia Mine Reclamation Symposium  in Cranbrook, BC, 1997. The Technical and Research Committee on Reclamation  40 Proceedings of the 21st Annual British Columbia Mine Reclamation Symposium  in Cranbrook, BC, 1997. The Technical and Research Committee on Reclamation  DISCUSSION Rhizobium Inoculation - Greenhouse and Mine Spoil Sporadic nodulation and poor legume growth is interpreted to indicate that the resident populations of indigenous  Rhizobia  in the mine spoil at the Line Creek mine are small. Only Oxytropis sericea occasionally possessed effective nodules. The number of nodules per plant was much lower than the numbers found on agronomic species such as Medicago sativa (Allen and Allen 1981). The small number of nodules may be an indication of a more ef r conservative nitrogen metabolism for these legume species. Location of the nodules was generally in the peripheral zone rather than on the root crown. This is comparable to most agronomic species (Johnson and Rumbaugh 1981). In general, there was a lower number of nodules and effective nodules per plant in the greenhouse experiment when compared to the mine spoil, a result observed by Johnson and Rumbaugh (1986). There did not appear to be any relationship between the number of nodules per plant and legume species. However, Astragalus alpinus is apparently more promiscuous than all the other species. All of the host plants appear to be sensitive to soil nitrogen levels, a physiological response which generally results in an increased number of ineffective nodules (Kenny and Cuany 1990).41 Proceedings of the 21st Annual British Columbia Mine Reclamation Symposium  in Cranbrook, BC, 1997. The Technical and Research Committee on Reclamation Mycorrhizal Relationships The fungal association results are in general agreement with that of Currah and van Dyk (1986). Most of the alpine species had extensive surface nets of dematiaceous septate fungi (DSF). Only Hedysarum sulphurescens was infected with vesicular arbuscular mycorrhizal. DSF infections of the Hedysarum species increased with an increase in elevation and exposure. The habitats of the alpine populations of all species were, in general, Regosols with small amounts of soil organic matter. The absence of VAM fungi in such soils has been documented in earlier research (V?re et al. 1992). Therefore, the present study confirms the results of previous studies as well as increases the number of known legume species which have either VAM or DSF associations. Mycorrhizal Inoculation - Mine Spoil The results of the fungal inoculation field experiment revealed that inoculation with either VAM or DSF fungi provides an advantage to the legume species. Survival, above ground biomass and reproductive activity were greater for the dual inoculation treatment of each species. Although yields were not quantified and detailed fertilizer trials were not conducted, it is expected that dual inoculation would result in the most favorable establishment results for these species. Inoculation with DSF and VAM fungi would also benefit the wide range of indigenous high elevation species that may be used in reclamation seed mixes. The results of this section confirm published results for other species (Azcon-Aguilar et al. 1982, Redente and Reeves 1981, Skujins and Allen 1986, Carpenter and Allen 1988, Miller and Jastrow 1992). The production and inoculation of Rhizobia does not appear to present a serious problem from a operational perspective. However, mycorrhizal inoculation will be more difficult because of the fungal culturing problems and inoculation difficulties (Call and McKeIl 1982). Nevertheless, attempts must be made to establish these microorganisms if successful establishment and survival of these legume species will be possible on high-elevation mine spoil. ACKNOWLEDGMENTS Funding for this project was provided by the Science Council of British Columbia, Fording Coal Limited -Fording River Operations and Line Creek Resources Limited.REFERENCES Allen, E.B. (1984). The role of mycorrhizae in mined land diversity. Third Biennial Symposium on Surface Coal Mine Reclamation on the Great Plains. (F.F. Munshower and S.E. Fisher, Jr., Co-chairmen) Reclamation Unit, Montana State University, Billings, pp. 273-295. Allen, E.B. and M.F. Allen. (1980). Natural re-establishment of vesicular-arbuscular mycorrhizae following strip-mine reclamation in Wyoming. Journal of Applied Ecology, 17, 139-147. Allen, E.B., J.C. Chambers, KF. Connor, M.F. Allen and RW. Brown.  (1987). Natural re-establishment of mycorrhizae in disturbed alpine ecosystems. Arctic and Alpine Research, 19, 1, 11-20. Allen, M.F. and C.F. Friese. (1992). Mycorrhizae and reclamation success: importance and measurement.  Evaluating Reclamation Success: Ecological Consideration. Proceedings of the Annual Meeting of the American Society for Surface Mining and Reclamation, April 23-26, 1990, Charleston, West Virginia. United States Department of Agriculture Forest Service General Technical Report NE-164. (J.C. Chambers and G.L. Wade, Editors). Northeastern Forest Experimental Station, Radnor, pp. 17-25. 42 Proceedings of the 21st Annual British Columbia Mine Reclamation Symposium  in Cranbrook, BC, 1997. The Technical and Research Committee on Reclamation Allen, O.N. and E.K. Allen. (1981). The Leguminosae: A Source Book of Characteristics, Uses, and Modulation. The University of Wisconsin Press, Madison, Wisconsin. 812 pp.  Azc?n-Aguilar, C. and J.M. Barea. (1992). Interactions between mycorrhizal fungi and other rhizosphere microorganisms. Mycorrhizal Functioning: An Integrative Plant-Fungal Process. (M.F. Allen, Editor). Chapman and Hall Publishers, London, pp. 163-198. Azc?n-Aguilar, C., J.M. Barea, R. Azc?n and J. Olivares. (1982). Effectiveness of Rhizobium and VA mycorrhiza in the introduction of Hedysarum coronarium in a new habitat. Agriculture and Environment, 7, 199-206. Barea, J.M., F. EI Atrach and R. Azc?n. (1989). Mycorrhiza and phosphate interactions as affecting plant development N2 fixation, N transfer and N uptake from soil in legume-grass mixtures by using a N15 dilution technique. Soil Biology and Biochemistry, 21,4, 581-589. Barea, J.M., and C. Azc?n-Aquilar. (1983). Mycorrhizas and their significance in nodulating nitrogen-fixing plants. Advanced Agronomy, 36, 1-54. Bell, M.A.M. and C.R. Smyth. (1988). Native Legume Species for High Elevation Revegetation in the Rockies of British Columbia. Prepared for British Columbia Science Council by the Biology Department of the University of Victoria, Research Contract Numbers 80RC-11, 80RC-13, and 80RC-15, Vancouver. 261 pp. Brockwell, J., A. Diatloff, R.J. Roughley and R.A. Date. (1982). Selection of Rhizobia  for inoculants. Nitrogen Fixation in Legumes. (J.M. Vincent, Editor). Academic Press, Sydney, pp. 173-191. Brown, R.W. and J.C. Chambers. (1990). Reclamation practices in high-mountain ecosystems. Proceedings - Symposium on Whitebark Pine Ecosystems: Ecology and Management of a High Mountain Resource. (W. C. Schmidt, Compiler). United States Department of Agriculture - Forest Service. Intel-mountain Research Station General Technical Report INT-270, Ogden, Utah. pp. 329-334.Call, GA. and C.M. McKell. (1982). Vesicular-arbuscular mycorrhizae - a natural revegetation strategy for disposed oil shale. Reclamation and Revegetation Research, 1, 337-347. Carpenter, A.T. and M.F. Allen. (1988). Responses of Hedysarum bor?ale Nutt. to mycorrhizas and Rhizobium: plant and soil nutrient changes in a disturbed.shrub steppe. New Phytologist, 109, 125-132. Chapin, F.S. III. (1983). Patterns of nutrient absorption and use by plants from natural and man-modified environments. Disturbance and Ecosystems. Ecological Studies 44, (H.A. Mooney and m. Godron, Editors). Springer-Verlag Publishers, New York. pp. 259-275.Copeland, L.O. and M.B. McDonald. (1985). Principles of Seed Science and Technology. Second Edition. Burgess Publishing Company, Minneapolis. 336 pp.Currah, R.S. and M. van Dyk. (1986). A survey of some perennial vascular plant species native to Alberta for occurrence of mycorrhizal fungi. Canadian Field Naturalist, 100, 3, 330-342. Errington, J.C. (1979). Revegetation Studies in the Peace River Coal Block, 1978. Province of British Columbia, Ministry of Energy, Mines and Petroleum Resources, Inspection and Engineering Division. Paper 1979-3. Victoria. 79 pp. Fl?eler, R. and A. Hasler. (1990). Native legumes of the Swiss Alps in high altitude revegetation research. Proceedings: High Altitude Revegetation Workshop No. 9. (W.R. Keammerer and J. Todd, 43 Proceedings of the 21st Annual British Columbia Mine Reclamation Symposium  in Cranbrook, BC, 1997. The Technical and Research Committee on Reclamation Editors). Colorado Water Resources Research Institute, Information Series No. 63, Colorado State University, Fort Collins, pg. 228. (Abstract).Garvin, S. and W.C. Lindemann. (1983). A new plant growth technique for the most-probable number estimation of Rhizobium. Soil Science Society of America Journal, 47, 604-606. Gibson, A.H. (1980). Methods for legumes in glasshouses and controlled environment cabinets. Methods for Evaluating Biological Nitrogen Fixation. (FJ. Bergersen, Editor). John Wiley and Sons, New York, pp. 139-184. Hayman, D.S. (1986). Mycorrhizae of nitrogen-fixing legumes. MIRCEN Journal, 2, 121-145. Hayman, D.S., E.J. Morris and RJ. Page. (1981). Methods for inoculating field crops with mycorrhizal fungi. Annals of Applied Biology, 99, 247-253. Hoagland, D.R and T.C. Broyer. (1936). General nature of the process of salt accumulation by roots with description of experimental methods. Plant Physiology, 11, 471-507. Johnson, D.A. and M.D. Rumbaugh. (1981). Nodulation and acetylene reduction by certain rangeland legume species under field conditions. Journal of Range Management, 34, 3, 178-181. Johnson, D.A. and M.D. Rumbaugh. (1986). Field nodulation and acetylene reduction activity of high-altitude legumes in the western United States. Arctic and Alpine Research, 18, 171-179. Kenny, S. T. and RL. Cuany. (1990). Nitrogen accumulation and acetylene reduction activity of native lupines on disturbed mountain sites in Colorado. Journal of Range Management, 43, 1, 49-51. Klebesadel, L.J. (1971). Native Alaskan legumes studied. Agroborealis, 3, 1, 9-11. Lambert, D.H. and H. Cole, Jr. (1980). Effects of mycorrhizae on establishment and performance of forage species in mine spoil. Agronomy Journal,, 72, 257-260. Little, T.M. and F. J. Hills. (1978). Agricultural Experimentation. Design and Analysis. John Wiley and Sons, New York. 350 pp.Miller, RM. and J.D. Jastrow. (1992). The application of VA mycorrhizae to ecosystem restoration and reclamation. Mycorrhizal Functioning: An Integrative Plant-Fungal Process. (M.F. Allen, Editor). Chapman and Hall Publishers, London, pp. 438-467.Moore-Landecker, E. (1982). Fundamentals of the Fungi. Second Edition. Prentice-Hall, Incorporated, Engelwood Cliffs. 252 pp. Mullen, RB. and S.K. Schmidt. (1993). Mycorrhizal infection, phosphorus uptake and phenology in Ranunculus adoneus: implications for the functioning of mycorrhizae in alpine systems. Oecologia (Berlin), 94, 229-234. Phillips, J.M. and D.S. Hayman. (1970). Improved procedures for clearing roots and staining parasitic vesicular-arbuscular mycorrhizal fungi for rapid assessment of infection. Transactions of the British Mycological Society, 55, 158-161. Pojar, J., K. Klinka and D.V. Meidinger. (1987). Biogeoclimatic ecosystem classification in British Columbia. Forest Ecology and Management, 22, 119-154. 44 Proceedings of the 21st Annual British Columbia Mine Reclamation Symposium  in Cranbrook, BC, 1997. The Technical and Research Committee on Reclamation Redente, E.F. and F.B. Reeves. (1981). Interactions between vesicular arbuscular mycorrhiza and Rhizobium and their effect on sweetvetch growth. Soil Science, 132, 6, 410-415. Reeves, F.B., D. Wagner, T. Moorman and J. Kiel. (1979). The role of endomycorrhizae in revegetation practices in the semi-arid west. I. A comparison of incidence of mycorrhizae in severely disturbed versus natural environments. American Journal of Botany, 66, 6-13. Schenck, N.C. (Editor). (1982). Methods and Principles of Mycorrhizal Research. The American Phytopathological Society, St. Paul. 244 pp. Skousen, J.G. (1986). The use of legumes in mined land reclamation. Greenlands, 16, 2, 35-37. Skujins, J. and M.F. Allen. (1986). Use of mycorrhizae for land rehabilitation. MIRCEN Journal of Applied Microbiology and Biotechnology, 2, 161-176. Smith, S. and D.J. Read. (1996). Mycorrhizal Symbiosis. Academic Press, San Diego. 492 pp. Smreciu, E.A. (1993). Native Legumes for Reclamation in Alberta. RRTAC 93-9. Prepared for the Alberta Land Conservation and Reclamation Council by Wild Rose Consulting Incorporated, Edmonton. 94pp. Smyth, CR. (1987). Seed Production, Dispersal, Pr?dation and Germination of Seven High Elevation Rocky Mountain Legume Species. M. Sc. Thesis, Department of Biology, versity of Victoria, Victoria. 169 pp. Somasegaran, P. and H.J. Hoben. (1994). Handbook for Rhizobia. Methods in Legume-Rhizobia Technology. Springer-Verlag Publishers, New York. 460 pp. St. John, T.V. (1990). Mycorrhizal inoculation of container stock for restoration of self-sufficient vegetation. Environmental Restoration. Science and Strategies for Restoring the Earth. (JJ. Berger, Editor). Island Press, Covelo. pp. 103-112. V?re, H., M. Vestberg and S. Eurola. (1992). Mycorrhiza and root-associated fungi in Spitzbergen. Mycorrhiza, 1, 93-104. Vincent, J.M. (1970). A Manual for the Practical Study of the Root-Nodule Bacteria. IBP Handbook No. 15, Blackwell Scientific Publications, Oxford. 164 pp.Waaland, M.E. and E.B. Allen. (1987). Relationships between VA mycorrhizal fungi and plant cover following surface mining in Wyoming. Journal of Range Management, 40, 3, 271-276. [455] Young, J.A. and C.G. Young. (1986). Collecting, Processing and Germinating Seeds of Western WildlandPlants. Timber Press, Portland. 236 pp. 45 

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