Controls of nitrogen limitation in tallgrass prairie

Summary The relationship between fire frequency and N limitation to foliage production in tallgrass prairie was studied with a series of fire and N addition experiments. Results indicated that fire history affected the magnitude of the vegetation response to fire and to N additions. Sites not burned for over 15 years averaged only a 9% increase in foliage biomass in response to N enrichment. In contrast, foliage production increased an average of 68% in response to N additions on annually burned sites, while infrequently burned sites, burned in the year of the study, averaged a 45% increase. These findings are consistent with reports indicating that reduced plant growth on unburned prairie is due to shading and lower soil temperatures, while foliage production on frequently burned areas is constrained by N availability. Infrequent burning of unfertilized prairie therefore results in a maximum production response in the year of burning relative to either annually burned or long-term unburned sites.Foliage biomass of tallgrass prairie is dominated by C₄ grasses; however, forb species exhibited stronger production responses to nitrogen additions than did the grasses. After four years of annual N additions, forb biomass exceeded that of grass biomass on unburned plots, and grasses exhibited a negative response to fertilizer, probably due to competition from the forbs. The dominant C₄ grasses may out-compete forbs under frequent fire conditions not only because they are better adapted to direct effects of burning, but because they can grow better under low available N regimes created by frequent fire.     

Soil invertebrate and plant responses to mowing and carbofuran application in a North American tallgrass prairie

Relationships between soil invertebrate populations and primary production of a tallgrass prairie were investigated using the insecticide-nematicide carbofuran and a range of mowing intensities to manipulate invertebrate densities and resource quantity and quality. The trophic composition of nematode populations was monitored through each of two growing seasons. Earthworm and macroarthropod densities and primary production were assessed at the end of the second season. Invertebrate densities were generally reduced in carbofuran-treated plots, although individual weights of surviving macroarth-ropod herbivores increased significantly (p<0.05). Carbofuran failed to affect estimates of above- or belowground plant biomass after two years of treatment. Changes in resource quantity and quality resulted in rapid responses by dominant invertebrate consumer populations. A 28% reduction in live root mass and a 24% increase in root detritus following two years of mowing was associated with a 54% decrease in herbivorous nematode densities, a 47% increase in microbivorous nematode densities, and a 41% increase in native earthworm biomass.     

Relationship of seed source and arbuscular mycorrhizal fungi inoculum type to growth and colonization of big bluestem (Andropogon gerardii)

Big bluestem (Andropogon gerardii) is a key grass of tallgrass prairies and is commonly included in restoration projects. In many cases, it has been found to benefit significantly from arbuscular mycorrhizal (AM) fungi, however results have varied under non-sterile soil conditions. This study investigated the effects of two types of AM fungi inoculum (commercial and prairie) on growth and root colonization of big bluestem from five different seed sources grown in non-sterile soils. Seed sources were collected from five remnant prairies in the Tallgrass Prairie Peninsula located in the Midwestern United States. Growth of big bluestem and root length colonized by AM fungi was highly variable among seed source treatments. Overall, percentage of root length colonized by AM fungi was positively correlated with the total dry weight of plants, and plants that received inoculum generally grew better than those that did not receive inoculum. Inoculum treatment affected both big bluestem growth and percentage root length colonized and there was an interaction between seed source and inoculum treatment relative to colonization. Root colonization responses were not significantly different between the prairie and commercial inoculum types, although there was a significant response between plants that received additional inoculum as opposed to no additional inoculum. Seed sources from Ohio and Illinois had the highest biomasses and greatest percentage of root length colonized while plants from Wisconsin and Missouri grew poorly and had low root colonization. These results demonstrate the importance of considering both seed source and inoculum type before the incorporation of AM inoculum to prairie restoration projects.     

Effects of fire,topography and year-to-year climatic variation on species composition in tallgrass prairie

Native unploughed tallgrass prairie from Konza Prairie, Kansas, USA is described with respect to plant species compositional changes over a five year period in response to fire and topography. The principal gradient of variation in the vegetation is related to time since burning. Species show an individualistic response in terms of relative abundance to this gradient. Both the percentage of and cover of C₄ species and all grasses decrease as the prairie remains unburnt. Forb and woody plant species numbers and abundance increase along this gradient. A secondary gradient of variation reflects topography (i.e. upland versus lowland soils). Upland soils support a higher species richness and diversity. Upland and lowland plant assemblages are distinct except on annually burnt prairie. The interaction between burning regime, topography and year-to-year climatic variation affects the relative abundance of the plant species differentially. The most dominant species overall, Andropogon gerardii, was affected only by year-to-year variation (i.e. climate). Its position at the top of the species abundance hierarchy was unaffected by burning regime or soil type. The other dominant species showed a suite of varying responses to these factors.Deceased May, 1986.     

Light utilization and competition between Echinacea purpurea, Panicum virgatum and Ratibida pinnata under greenhouse and field conditions

Competitive interactions were compared under field and greenhouse conditions for three representative tallgrass prairie species, Echinacea purpurea, Panicum virgatum and Ratibida pinnata. These were planted in monoculture and in mixtures of two species using a replacement series design with groups of four or eight plants. Competition was determined from shoot dry weight data collected during 120days in the greenhouse and after 415days in the field. Yields declined with increased density of a single species in the greenhouse from 40days onward and in the field. Relative yields were up to 100% higher in mixtures than in monocultures for all species early on in the greenhouse experiment. Later in the experiment and in the field relative yields of E.purpurea decreased in the presence of P.virgatum and R.pinnata, whereas relative yields of these two species increased in the presence of E.purpurea. There were correlations in relative yield between the field and the greenhouse experiment at 80 and 120days. In the greenhouse P.virgatum maintained higher net assimilation rates than the other species. Relative growth rates of all species were higher in monoculture and in mixtures up to 40days, after which they declined, especially for E.purpurea in mixtures. In the field, higher light intensities occurred in pure stands of E.purpurea than in mixed stands with other species. The order of competitive ability that was apparent from these field and greenhouse studies, P.virgatum=R.pinnata>E.purpurea, could be partially explained by photosynthetic rates in relation to canopy light interception.     

Physiological Performance of Andropogon gerardii, Panicum virgatum, and Sorghastrum nutans on Reclaimed Mine Spoil

Physiological and vegetative performances of three prairie grasses were investigated to assess their adaptation to soil conditions at two strip mine sites and a nearby railroad prairie. Additionally, rhizomes of the species were transplanted to a pot experiment and grown in both field soil and greenhouse potting medium to investigate the extent to which plants are limited under field conditions. Field measurements of photosynthetic rate and stomatal conductance to water vapor were made on the three species monthly from May to late August. Gas exchange measurements on potted plants were made biweekly from early May to mid-July. In September, vegetative and flowering characteristics were measured on both field and potted plants. Field gas exchange rates were highest at one of the mines. Sorghastrum nutans had the highest rates at the mine sites, whereas Panicum virgatum had the highest rates at the prairie site. Potted plants from the prairie site usually exhibited the highest gas exchange rates, and Sorghastrum nutans had higher rates than Panicum virgatum and Andropogon gerardii. Potted plants in field soil generally had higher gas-exchange rates than plants growing in greenhouse potting medium, and potted plants had higher gas-exchange rates than field-grown plants. Vegetative and reproductive performance of field plants was highest at one of the mine sites. Potted plants in greenhouse medium had up to twice the vegetative and reproductive output as potted plants in field soil or plants growing in the field. The physiological and vegetative performance of these species indicates that they are well adapted to the soil conditions at these strip mine sites, and that they are a viable alternative to nonnative plantings for restoration.     

Effects of elevated CO2 and nitrogen fertilization pretreatments on decomposition on tallgrass prairie leaf litter

Standing dead and green foliage litter was collected in early November 1990 from Andropogon gerardii (C₄), Sorghastrum nutans (C₄), and Poa pratensis (C₃) plants that were grown in large open-top chambers under ambient or twice ambient CO₂ and with or without nitrogen fertilization (45 kg N ha⁻¹). The litter was placed in mesh bags on the soil surface of pristine prairie adjacent to the growth treatment plots and allowed to decay under natural conditions. Litter bags were retrieved at fixed intervals and litter was analyzed for mass loss, carbon chemistry, and total Kjeldahl nitrogen and phosphorus. The results indicate that growth treatments had a relatively minor effect on the initial chemical composition of the litter and its subsequent rate of decay or chemical composition. This suggests that a large indirect effect of CO₂ on surface litter decomposition in the tallgrass prairie would not occur by way of changes in chemistry of leaf litter. However, there was a large difference in characteristics of leaf litter decomposition among the species. Poa leaf litter had a different initial chemistry and decayed more rapidly than C₄ grasses. We conclude that an indirect effect of CO₂ on decomposition and nutrient cycling could occur if CO₂ induces changes in the relative aboveground biomass of the prairie species.     

Comparative physiology of burned and unburned Rhus laurina after chaparral wildfire

Summary Laurel Sumac (Rhus laurina) is a dominant member of the coastal chaparral community of southern California that survives periodic burning by wildfires by resprouting from a lignotuber (root crown). We investigated the physiological basis for resprouting by comparing shoot elongation, leaf nitrogen content, tissue water status, leaf conductance to water vapor diffusion, and photosynthetic rates of post-fire R. laurina to those of adjacent unburned shrubs. Resprouts had higher rates of shoot elongation, leaf conductance, and photosynthesis than mature, unburned shrubs. Leaf nitrogen contents were elevated in burned shrubs even though their leaves developed interveinal chlorosis. A comparison of soil water potential to predawn water potential indicated that roots of R. laurina remain active below 2 m during the first summer drought after wildfire. Our results support the hypothesis that lignotubers not only contain dormant buds that develop into aerial shoots after wildfire but they also supply nutrient resources that enhance shoot elongation. Because R. laurina is relatively sensitive to drought, yet very successful in its rapid recovery after fire, maintaining an active root system after shoot removal may be the primary function of the massive lignotuber formed by this species.     

Effects of collembola (arthropoda) and relative germination date on competition between mycorrhizalPanicum virgatum (Poaceae) and non-mycorrhizalBrassica nigra (Brassicaceae)

In this experiment the separate and interactive effects of grazing of vesicular-arbuscular mycorrhizal hyphae by collembola and of relative germination date on competition between the mycorrhizal perennial grassPanicum virgatum and the non-mycorrhizal annual cruciferBrassica nigra were investigated. In the absence of competition,P. virgatum mass and P uptake were not affected by collembola grazing; grazing did reduce tissue N concentration and root: shoot ratio. Competition fromB. nigra plants of the same age/size (“simultaneous competition”) significantly reducedP. virgatum total, root, and shoot mass relative to control plants not subject to competition. In contrast, when in competition,B. nigra plants did not differ in biomass fromB. nigra controls grown without competition. Simultaneous competition also reduced N and P uptake byP. virgatum, but not byB. nigra. Grazing by collembola during simultaneous competition increased the differences in nutrient uptake and tended to shift the competitive balance further toward the non-mycorrhizalB. nigra. WhenP. virgatum plants were subjected to competition fromB. nigra plants which germinated three weeks later (“offset competition”) the situation was reversed: offsetB. nigra plants were negatively affected by competition while the larger, olderP. virgatum plants were not. Thus, relative germination date is important in determining the relative competitive ability of these two species. Grazing by collembola did not affect offset competition. The grazing of VAM hyphae by collembola appears to increase N availability in this experimental system. Under simultaneous competition, this N is taken up by the more extensive root system ofB. nigra; under offset conditions, the root system of the smallerB. nigra plants is insufficient to take advantage of the added resources. Thus, we suggest that grazing-induced transient changes in nutrient availability and the differential abilities of the two species to make use of these added resources constitute the mechanisms by which relative germination date and collembola grazing influence competition.     

Production on burned and unburned sand prairies during drought and non-drought years

Biomass production was determined on burned and unburned sand prairie sites in Illinois during the 1986–89 growing seasons. Severe droughts characterized the 1988 and 1989 growing seasons with precipitation being at least 50% less in these two years than in 1985 and 1986. During the growing season following a fire, in drought and non-drought years, burned sites had higher total green biomass (all species), little bluestem (Schizachyrium scoparium) root and shoot biomass, and total grass and forb biomass production than the unburned sites. In 1988 and 1989, production on all sites was frequently significantly lower each month than it was in 1986 or 1987. The differences in productivity between burn and unburned sites were less pronounced during the drought years (1988–1989) than during the non-drought years (1986–87). Grass productivity (all species) declined considerably during drought years. In contrast, forb productivity did not vary significantly among years. Despite the drought, productivity of all species and little bluestem (root and shoot combined) was best predicted by the burn condition of the sites. Unlike many studies where drought alone can have significant effects on productivity, this study shows that on sand prairies the burn condition of sites, or burning regimes, and associated influences, can have a greater substantial impact on productivity than drought. Moisture deficits due to drought may, however, have longer term effects than those detected during the span of this study.     

Effects of Switchgrass (Panicum virgatum) Rotations with Peanut (Arachis hypogaea L.) on Nematode Populations and Soil Microflora

A 3-year field rotation study was conducted to assess the potential of switchgrass (Panicum virgatum) to suppress root-knot nematodes (Meloidogyne arenaria), southern blight (Sclerotium rolfsii), and aflatoxigenic fungi (Aspergillus sp.) in peanut (Arachis hypogaea L.) and to assess shifts in microbial populations following crop rotation. Switchgrass did not support populations of root-knot nematodes but supported high populations of nonparasitic nematodes. Peanut with no nematicide applied and following 2 years of switchgrass had the same nematode populations as continuous peanut plus nematicide. Neither previous crop nor nematicide significantly reduced the incidence of pods infected with Aspergillus. However, pod invasion by A. flavus was highest in plots previously planted with peanut and not treated with nematicide. Peanut with nematicide applied at planting following 2 years of switchgrass had significantly less incidence of southern blight than either continuous peanut without nematicide application or peanut without nematicide following 2 years of cotton. Peanut yield did not differ among rotations in either sample year. Effects of crop rotation on the microbial community structure associated with peanut were examined using indices for diversity, richness, and similarity derived from culture-based analyses. Continuous peanut supported a distinctly different rhizosphere bacterial microflora compared to peanut following 1 year of switchgrass, or continuous switchgrass. Richness and diversity indices for continuous peanut rhizosphere and geocarposphere were not consistently different from peanut following switchgrass, but always differed in the specific genera present. These shifts in community structure were associated with changes in parasitic nematode populations.     

The effect of CO2 enrichment on leaf photosynthetic rates and instantaneous water use efficiency of Andropogon gerardii in the tallgrass prairie

Open-top chambers were used to study the effects of CO₂ enrichment on leaf-level photosynthetic rates of the C₄ grass Andropogon gerardii in the native tallgrass prairie ecosystem near Manhattan, Kansas. Measurements were made during a year with abundant rainfall (1993) and a year with below-normal rainfall (1994). Treatments included: No chamber, ambient CO₂ (A); chamber with ambient CO₂ (CA); and chamber with twice-ambient CO₂ (CE). Measurements of photosynthesis were made at 2-hour intervals, or at midday, on cloudless days throughout the growing season using an open-flow gas-exchange system. No significant differences in midday rates of photosynthesis or in daily carbon accumulation as a result of CO₂ enrichment were found in the year with abundant precipitation. In the dry year, midday rates of photosynthesis were significantly higher in the CE treatment than in the CA or A treatments throughout the season. Estimates of daily carbon accumulation also indicated that CO₂ enrichment allowed plants to maximize carbon acquisition on a diurnal basis. The increased carbon accumulation was accounted for by greater rates of photosynthesis in the CE plots during midday. During the wet year, CO₂ enrichment decreased stomatal conductance, which allowed plants to decrease transpiration while still photosynthesizing at rates similar to plants in ambient conditions. During the dry year, CO₂ enrichment allowed plants to maintain photosynthetic rates even though stomatal conductance and transpiration had been reduced in all treatments due to stress. Estimates of instantaneous water-use efficiency were reduced under CO₂ enrichment for both years. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of grazing intensity on growth,reproduction, and abundance of three palatable forbs in Kansas tallgrass prairie

The effects of different intensities of cattle grazing on theaboveground growth, reproduction, and abundances of three palatable forbs werestudied in native tallgrass prairie. Populations of Asterericoides, Ruellia humilis, andAmorpha canescens were sampled at peak flowering duringthe1993–1995 growing seasons in four annually-burned sites varying incattle stocking density [ungrazed, low, moderate, high]. The threeforbs exhibited reduced shoot growth and/or reproduction under moderate toheavy grazing, and in no case did grazing increase any measure of plantperformance. Ruellia showed reduced shoot height andbiomass, percentage of stems flowering, and reproductive biomass in response tograzing. Aster showed decreases in shoot biomass andheightwith grazing. Amorpha showed no change in shoot orreproductive biomass, but a decrease in percent of flowering stems and inreproductive allocation with grazing. Patterns in the percentage of stemsgrazedindicated generally high but variable palatability among these species. Bycontrast, the three species showed inconsistent population response to grazing.Abundance (frequency) of all three species indicated no short-term changebetween years in response to grazing intensity. Responses of these speciesdiffer considerably from those of most other perennial tallgrass prairie forbsthat are unpalatable, unconsumed, and increase in performance (e.g. size,abundance) due to release from competition from the dominant grasses underungulate grazing. The results demonstrate that immediate aboveground growth andreproductive responses of established adults to grazing are not good predictorsof grazer effects on population abundances in tallgrass prairie.     

Ultrastructural changes in suspension culture cells of Panicum maximum during cryopreservation

A Panicum maximum cell suspension was used to study ultrastructural changes during cryopreservation. Pregrowing the cells in mannitol caused reduction in the vacuolar volume by redistribution of the large central vacuole into a number of smaller vesicles. Invaginations were formed in the plasma membrane of the cells, to accommodate the reduced cell volume. Swelling of organelles occurred during different stages of cryopreservation. The cisternae of the endoplasmic reticulum dilated and formed vesicles. Although some damage was apparent, organelles were still recognizable in cells frozen slowly and freeze-fixed at –10°C. The cells were able to repair such damage within two days in culture, and regained their normal appearance. Cells frozen slowly without any cryoprotection, and cells frozen rapidly by direct immersion into liquid nitrogen after cryoprotection, were lethally damaged by destruction of membranous structures. Osmiophilic granules were found along the plasma membrane of lethally damaged cells, indicating that their formation is a consequence of freeze damage, rather than a mechanism to prevent injury.Abbreviations 2,4-D 2,4-dichlorophenoxyacetic acid - DMSO dimethyl sulfoxide     

Effects of seasonal fire,bison grazing and climatic variation on tallgrass prairie vegetation

We used univariate and multivariate techniques to evaluate vegetation-environment relationships for plant functional groups on a tallgrass prairie site in northern Oklahoma, USA burned seasonally and grazed by bison (Bison bison L.). The objective of the study was to identify important environmental variables associated with variation in residual aboveground standing crop (phytomass) and abundance of plant functional groups. Phytomass was predictably linked to season, with highest levels found in the latter portion of the growing season when the warm-season tallgrasses that dominated the site were most actively growing. When the effects of seasonal phenology were removed, stepwise regression revealed that phytomass variation was best explained by year-to-year climatic variation, seasonal burn type, and bison grazing. Phytomass was negatively related to bison grazing under all conditions. A number of plant functional groups responded to individual environmental variables: relative abundance levels of tallgrasses, little bluestem, annual grasses, forbs and legumes all varied with burn season; little bluestem, annual grasses, and sedges varied by topoedaphic position, while forbs exhibited a positive relationship with bison grazing intensity. Canonical Correspondence Analysis (CCA) was used to directly relate composition patterns of functional groups to environmental variables. CCA bi-plot of the ordination revealed that annual grasses were most closely associated with summer fires; sedges were associated with fall burns and the 1995 sampling year; legumes associated with a gradient representing the length of time since fire; while tallgrasses and little bluestem ordinated nearest a gradient representing bison grazing. Forbs and perennial grasses did not clearly associate with any particular environmental gradient, suggesting they were either simultaneously affected by several parameters or that environmental attributes important to these groups were not measured. Total phytomass and sedges were strongly influenced by yearly climatic variation. Relative abundance of some plant functional groups was principally determined by a single factor, while others were equally influenced by a suite of environmental interactions.     

Anti-tick effects of Melinis minutiflora and Andropogon gayanus grasses on plots experimentally infested with Boophilus microplus larvae

Our objective in this trial was to investigate the anti-tick effects of molasses grass (Melinis minutiflora) and gamba grass (Andropogon gayanus) against Boophilus microplus, using buffel grass (Cenchrus ciliaris) as a control. During a 3-year-period fieldwork was carried out in the tropical, subhumid climate at Progreso, Municipality of Jiutepec, Morelos, Mexico. Experimental plots were established for three treatments with six replicates under a random factorial design in order to evaluate each of the four seasons. Additional work was developed for two more consecutive autumn seasons to evaluate the anti-tick effects of these grasses. Once the plots were established they were infested with 5000 B. microplus larvae and sampled by dragging flannel cloths. The anti-tick effects were measured by counting the number of larvae recovered from each of the experimental plots. Both grasses, M. minutiflora and A. gayanus, demonstrated anti-tick effects; however, M. minutiflora grass had the greatest anti-tick effects through the seasons studied and in the three consecutive autumns evaluated.     

Flowering Phenology and Genetic Similarity among Local and Recently Introduced Populations of Andropogon gerardii in Ohio

In Ohio and elsewhere, recent grassland plantings in the federal Conservation Reserve Program (CRP) have become much more extensive than native prairie remnants. The seed source for CRP grasslands in Ohio often comes from as far away as Missouri or Texas, which may be undesirable from the standpoint of conservation genetics. The goal of this study was to examine the potential for gene flow from large, recently introduced populations of Big bluestem (Andropogon gerardii, Poaceae) to small local populations of this outcrossing perennial species. We examined the potential for cross‐pollination between three local populations and three introduced CRP populations by comparing flowering phenologies. Flowering times overlapped extensively, indicating that cross‐pollination is possible where local and introduced genotypes co‐occur. To compare genetic variation in local and CRP populations, we analyzed variation at 68 RAPD loci in six populations of each type. Somewhat surprisingly, we found no significant differences in the genetic diversity or composition between the two groups (local vs. CRP). In summary, we found that local and introduced populations of Big bluestem have the potential to interbreed, based on their flowering periods, but further research is needed to determine whether local genotypes harbor unique genetic variation that could be jeopardized by hybridization with introduced genotypes.