The timing and degree of root proliferation in fertile-soil microsites for three cold-desert perennials

Summary Root proliferation in nutrient-rich soil patches is an important mechanism facilitating nutrient capture by plants. Although the phenomenon of root proliferation is well documented, the specific timing of this proliferation has not been investigated. We studied the timing and degree of root proliferation for three perennial species common to the Great Basin region of North America: a shrub, Artemisia tridentata, a native tussock grass, Agropyron spicatum, and an introduced tussock grass, Agropyron desertorum. One day after we applied nutrient solution to small soil patches, the mean relative growth rate of Agropyron desertorum roots in these soil patches was two to four times greater than for roots of the same plants in soil patches reated with distilled water. Most of the increased root growth came from thin, laterally branching roots within the patches. This rapid and striking root proliferation by Agropyron desertorum occurred in response to N-P-K enrichment as well as to P or N enrichment alone. A less competitive bunchgrass, Agrophyron spicatum, showed no tendency to proliferate roots in enriched soil patches during these two-week experiments. The shrub Artemisia tridentata proliferated roots within one day of initial solution injection in the N-enrichment experiment, but root proliferation of this species was more gradual and less consistent in the N-P-K and P-enrichment experiments, respectively. The ability of Agropyron desertorum to proliferate roots rapidly may partly explain both its general competitive success and its superior ability to exploit soil nutrients compared to Agropyron spicatum in Great Basin rangelands of North America.     

Comparative demography of co-occurring introduced and native tussock grasses: persistence and potential expansion

Summary Demographic characteristics associated with the maintenance and growth of populations, such as seed dynamics, seedling emergence, survival, and tiller dynamics were examined for two tussock grasses, the native Agropyron spicatum and the introduced Agropyron desertorum in a 30-month field study. The introduced grass was aerially sown onto a native grassland site. Seed production of the introduced grass was greater than the native grass in both above- and below-average precipitation years. Seeds of A. spicatum were dispersed when they mature, while A. desertorum retained some seeds in inflorescences, and dispersed them slowly throughout the year. This seed retention allowed some seeds of the introduced grass to escape peak periods of seed predation during the summer and allowed seeds to be deposited constantly into the seed bank. Carryover of seeds in the seed bank beyond one year occurred in the introduced grass but not in the native species. For both species, seedling emergence occurred in both autumn or spring. Survival rates for A. desertorum were higher than A. spicatum when seedlings emerged between November and March. Survival rates of cohorts emerging before November favored A. spicatum whereas survival rates did not differ between species for cohorts emerging after March. Individuals of both species emerging after April were unable to survive the summer drought. Demographic factors associated with seeds of A. desertorum seemed to favor the maintenance and spread of this introduced grass into native stands formerly dominated by A. spicatum.     

Characteristics of successful competitors: an evaluation of potential growth rate in two cold desert tussock grasses

Summary Within the first few weeks after seedling emergence, Agropyron desertorum, a more competitive tussock grass, had a much higher mean relative growth rate (RGR) than Agropyron spicatum, a very similar, but less competitive species. However, beyond the early seedling stage, the two grasses had a remarkably similar whole-plant RGR in hydroponic culture and aboveground RGR in glasshouse soil, if root temperatures were above approximately 12°C. At soil temperatures between 5 and 12°C, A. desertorum exhibited a 66% greater aboveground RGR than A. spicatum (P<0.05). Both species responded similarly to warming soil temperatures. In the field, however, tiller growth rates were generally similar. Neither species showed marked tiller elongation until a couple of weeks after snowmelt, by which time soil temperatures, at least to a depth of 10 cm, were above 12°C for a significant portion of the day. Aboveground biomass accumulation over a three-year period indicated that both grasses had similar potential growth rates whereas Artemisia tridentata ssp. vaseyana, a common neighbor planted in the same plots, had a much greater potential growth rate. The greater competitive ability of adult A. desertorum, as compared to A. spicatum, cannot be attributed to appreciable differences in potential growth rates.     

Coping with herbivory: Photosynthetic capacity and resource allocation in two semiarid Agropyron bunchgrasses

Summary Agropyron desertorum, a grazing-tolerant bunchgrass introduced to the western U.S. from Eurasia, and Agropyron spicatum, a grazing-sensitive bunchgrass native to North America, were examined in the field for photosynthetic capacity, growth, resource allocation, and tiller dynamics. These observations allowed identification of physiological characteristics that may contribute to grazing tolerance in semiarid environments. A uniform matrix of sagebrush, Artemisia tridentata, provided an ecologically relevant competitive environment for both bunch-grass species. Physiological activity, growth, and allocation were also followed during recovery from a severe defoliation treatment and were correlated with tiller dynamics.Potential photosynthetic carbon uptake of both species was dominated by stems and leaf sheaths during June, when maximum uptake rates occurred. For both species, water use efficiency of stems and sheaths was similar to that of leaf blades, but nitrogen investment per photosynthetic surface area was less than in blades. In addition, soluble carbohydrates in stems and sheaths of both species constituted the major labile carbon pools in control plants. Contrary to current theory, these findings suggest that culms from which leaf blades have been removed should be of considerable value to defoliated bunchgrasses, and in the case of partial defoliation could provide important supplies of organic nutrients for regrowth. These interpretations, based on total pool sizes, differ markedly from previous interpretations based on carbohydrate concentrations alone, which suggested that crowns contain large carbohydrate reserves. In this study, crowns of both species contained a minor component of the total plant carbohydrate pool.Following defoliation, A. desertorum plants rapidly reestablished a canopy with 3 to 5 times the photosynthetic surface of A. spicatum plants. This difference was primarily due to the greater number of quickly growing new tillers produced following defoliation. Agropyron spicatum produced few new tillers following defoliation despite adequate moisture, and carbohydrate pools that were equivalent to those in A. desertorum.Leaf blades of regrowing tillers had higher photosynthetic capacity than blades on unclipped plants of both species, but the relative increase, considered on a unit mass, area, or nitrogen basis, was greater for A. desertorum than for A. spicatum. Agropyron desertorum also had lower investment of nitrogen and biomass per unit area of photosynthetic tissues, more tillers and leaves per bunch, and shorter lived stems, all of which can contribute to greater tolerance of partial defoliation.Greater flexibility of resource allocation following defoliation was demonstrated by A. desertorum for both nitrogen and carbohydrates. Relatively more allocation to the shoot system and curtailed root growth in A. desertorum resulted in more rapid approach to the preclipping balance between the root and shoot systems, whereas root growth in A. spicatum continued unabated following defoliation. Nitrogen required for regrowth in both species was apparently supplied by uptake rather than reserve depletion. Carbohydrate pools in the shoot system of both species remained very low following severe defoliation and were approximately equivalent to carbon fixed in one day by photosynthesis of the whole canopy.Dedicated to Drs. Michael Evenari and Konrad Springer     

Root growth response to defoliation in two Agropyron bunchgrasses: field observations with an improved root periscope

Summary Root growth responses to defoliation were observed in the field with an improved root periscope technique, which is described. The grazing tolerant, Eurasian bunchgrass, Agropyron desertorum, was compared with the very similar but grazing sensitive, North American bunchgrass, A. spicatum. Root length growth of clipped A. desertorum was about 50% of that of intact plants, while root elongation of clipped A. spicatum continued relatively unabated during ninety days of regrowth following severe defoliation. The reduced root growth in A. desertorum was correlated with the allocation of relatively more resources to aboveground regrowth, thus aiding reestablishment of the root: shoot balance. This balance was apparent in similar root mortality patterns of clipped and control A. desertorum plants in the season following defoliation. In clipped A. spicatum, however, root mortality increased in the winter following the season in which the clipping was done and continued into the subsequent growing season. Reduction of root growth following defoliation appears to be an effective mechanism to aid reestablishment of the photosynthetic canopy and the root: shoot balance. As such it contributes to both herbivory tolerance and maintenance of competitive ability.     

Phytoremediation Effect and Growth Responses of Cynodon spp. and Agropyron desertorum in a Petroleum-Contaminated Soil

In order to compare the petroleum tolerance and phytoremediation ability of a native grass, Agropyron desertorum (desert Wheatgrass) with Cynodon spp. (Bermuda grass) in a petroleum hydrocarbon-contaminated soil, a 7-month greenhouse experiment was performed. There were 4 soil treatments with 0% (uncontaminated soil), 2%, 4%, and 12% (w_oil/w_soil) petroleum concentration. Parameters including shoot and root fresh weight and dry weight, root penetration depth and root density depth, soil respiration, and total petroleum hydrocarbons (TPH) degradation were measured during and after experiments. The results showed an increase in shoot fresh weight of A. desertorum in soil polluted with 2% petroleum sludge compared to the uncontaminated soil, whereas the growth of Bermuda grass significantly decreased in corresponding treatment. Root growth of A. desertorum was decreased in 2% and 4% petroleum sludge, whereas it was increased in Bermuda grass species. Overall, root fresh weight of Bermuda grass was higher than that of A. desertorum in all treatments. Significant increase in microorganisms' activity was observed in the presence of petroleum sludge and plants in soil compared with uncontaminated soil without plants, and the highest soil respiration (37.6 mg C-CO₂/kg soil day) has been observed in the rhizosphere of Bermuda grass in treatment with 12% petroleum sludge. Plants had a significant role in the degradation of soil contaminants as TPH degradation in planted soils was significantly higher than that in unplanted soil (TPH degradation (%) was 30.4 and 38.9 in A. desertorum and Bermuda grass, respectively, whereas it was just 13.3 in unplanted soil). The rhizosphere of Bermuda grass had significantly less residual TPHs compared to A. desertorum. The results indicated that both Cynodon spp. and A. desertorum had a peculiar tolerance to petroleum pollution. Therefore, as Bermuda grass has already been suggested to be a typical and efficient species for phytoremediating petroleum-contaminated sites, A. desertorum may also prove to be a suitable native alternative.     

Inorganic N turnover and availability in annual- and perennial-dominated soils in a northern Utah shrub-steppe ecosystem

The exotic annual grass Bromus tectorum has replaced thousands of hectares of native perennial vegetation in semi-arid ecosystems of the western United States. Inorganic N availability and production were compared in soil from monodominant patches of Bromus tectorum, the perennial bunchgrass Elymus elymoides, and the shrub Artemisia tridentata, in Curlew Valley, a salt-desert shrub site in Northern Utah. Bromus-dominated soil had greater %N in the top 10 cm than Artemisia or Elymus-dominated soils. As determined by spring isotope-dilution assays, gross mineralization and nitrification rates were higher in Bromus-dominated than Artemisia-dominated soils, but gross rates of NH₄ ⁺ and NO₃ ^– consumption were also higher. Litterbags had greater mass loss and N mineralization when buried in Bromus stands than in Artemisia stands, indicating the soil environment under the annual grass promotes decomposition. As determined by nitrification potential assays, nitrifier populations were higher under Bromus than under Artemisia and Elymus. Soil inorganic N concentrations were similar among vegetation types in the spring, but NO₃ ^– accumulated under Bromus once it had senesced. An in situ net mineralization assay conducted in autumn indicated that germinating Bromus seedlings are a strong sink for soil NO₃ ^–, and that net nitrification is inherently low in soils under Artemisia and Elymus. Results of the study suggest that differences in plant uptake and the soil environment promote greater inorganic N availability under Bromus than under perennial species at the site.     

Effect of competition on stable carbon isotope ratios of two tussock grass species

Summary Previous studies have shown that plant carbon isotope composition varies when plants experience differences in water and nutrient availability. However, none have addressed the effect of root interactions, including competition for these soil resources, on carbon isotope ratios. We studied the effect of interspecific root interactions on the productivity and carbon isotope ratios of two Great Basin tussock grass species (Agropyron desertorum and Pseudoroegneria spicata). We compared grasses grown in mixture with sagebrush (Artemisia tridentara) to grasses in similar mixtures but where root interactions with sagebrush were limited by fiberglass partitions. During both years of the study, tussocks growing in competition with sagebrush produced tissue with more negative ¹³C values than grasses experiencing limited root interaction with sagebrush. The magnitude of this difference (0.5 to 0.9%) is similar to that found in other studies when soil fertility and moisture availability were altered.     

Species interactions at the level of fine roots in the field: influence of soil nutrient heterogeneity and plant size

Interference at the level of fine roots in the field was studied by detailed examination of fine root distribution in small soil patches. To capture roots as they occur in natural three-dimensional soil space, we used a freezing and slicing technique for microscale root mapping. The location of individual roots intersecting a sliced soil core surface was digitized and the identity of shrub and grass roots was established by a chemical technique. Soil patches were created midway between the shrub, Artemisia tridentata, and one of two tussock grasses, Pseudoroegneria spicata or Agropyron desertorum. Some soil patches were enriched with nutrients and others given only deionized water (control); in addition, patches were located between plants of different size combination (large shrubs with small tussock grasses and small shrubs with large tussock grasses). The abundance of shrub and grass roots sharing soil patches and the inter-root distances of individual fine roots were measured. Total average rooting density in patches varied among these different treatment combinations by only a factor of 2, but the proportion of shrub and grass roots in the patches varied sixfold. For the shrub, the species of grass roots sharing the patches had a pronounced influence on shrub root density; shrub roots were more abundant if the patch was shared with Pseudoroegneria roots than if shared with Agropyron roots. The relative size of plants whose roots shared the soil patches also influenced the proportion of shrub and grass roots; larger plants were able to place more roots in the patches than were the smaller plants. In the nutrient-enriched patches, these influences of grass species and size combination were amplified. At the millimeter- to centimeter-scale within patches, shrub and grass roots tended to segregate, i.e., avoid each other, based on nearest-neighbor distances. At this scale, there was no indication that the species-specific interactions were the result of resource competition, since there were no obvious patterns between the proportion of shrub and grass roots of the two species combinations with microsite nutrient concentrations. Other potential mechanisms are discussed. Interference at the fine-root level, and its species-specific character, is likely an influential component of competitive success, but one that is not easily assessed.     

Impact of early root competition on fitness components of four semiarid species

Summary Plant demographic and root exclusion approaches were used to examine the influence of roots of adult Artemisia tridentata, Agropyron desertorum, and Agropyron spicatum individuals on seedling survival of four C₃ semiarid species, three perennials, Ar. tridentata, Ag. desertorum, Ag. spicatum, and an annual, Bromus tectorum. Furthermore, height of Ar. tridentata seedlings and seed production of B. tectorum were assessed. The probability of a seedling being alive significantly depended on the seedling species, the neighboring adult species, and on the depth to which root competition was excluded. As seedlings, survival of Agropyron species did not differ, whereas survival of Ar. tridentata seedlings was higher than Ag. desertorum and was similar to Ag. spicatum. Bromus tectorum maintained significantly higher survival rates than perennial seedlings. Established individuals of Ar. tridentata reduced seedling survival more than established individuals of either Agropyron species. Seedling survival significantly increased with greater depth of root exclusion for the perennials but did not significantly affect seedling survival of B. tectorum. Height of Ar. tridentata seedlings and seed production of B. tectorum significantly increased with depth of root exclusion. Seed production of B. tectorum was highest when competing with Ag. desertorum and was lowest with Ar. tridentata. Root competition decreased the seed population of B. tectorum in the next generation even though it had no impact on survival. Competition in the upper soil horizon occurs between seedlings and established adults early in the growing season and potentially restricts root growth of seedlings. In arid and semiarid ecosystems, soil moisture is depleted from the upper horizons first, resulting in the death of seedlings that do not have access to moisture.     

Light field heterogeneity among tussock grasses: Theoretical considerations of light harvesting and seedling establishment in tussocks and uniform tiller distributions

Although the tussock growth form of caespitose graminoids is widespread, the effect of this growth form on light interception and carbon gain of tillers has received little attention. Daily incident photosynthetic photon flux density (PFD_inc) and carbon gain in monospecific stands of tussock grasses were compared with those of a hypothetical distribution with the equivalent tiller density per total ground area, but evenly distributed rather than clumped in tussocks. This was computed for two tussock grasses Pseudoroegneria spicata (Pursh) A. Löve (bluebunch wheatgrass) and Agropyron desertorum (Fisch, ex Link) Schult. (creasted wheatgrass) at different plant densities. Daily PFD_inc and net photosynthesis (A) were greater if tillers were distributed uniformly rather than clumped in tussocks, except when the density of tussocks was so great as to approach a uniform canopy. When tussock density per ground area was low, much of the difference between tussock and uniform tiller densities in PFD_inc and A was due to shading within the tussocks; up to 50–60% of the potential carbon gain was lost in A. desertorum due to shading within tussocks. In a matrix of tussocks, the light field for establishing seedlings was very heterogeneous; potential A ranged from 7 to 96% relative to an isolated seedling. The mean of daily PFD_inc and A for seedlings in a tussock stand were nearly identical to the values in corresponding stands of uniform tiller distributions. It is hypothesized that the loss of A resulting from clumping tillers into tussocks is offset by benefits of protecting sequestered belowground resources from invasion by seedlings of competitors.     

Distribution of VA mycorrhiza on halophytes on inland salt playas

The value of mycorrhizal association for higher plants has been well established. However, the impact of high salinity on the mycorrhizal relationship has not been investigated to any great extent. Inland salt playas represent an opportunity to test the impact of salinity because it is possible to obtain a gradient by following a transect from the centre of the salt playa to the higher outer zones. In a salt playa near Goshen, Utah, the sodium concentration ranged from 27,150 ppm in the centre to 25 ppm in the outer zone. In the playas with sodium concentrations of 20,000 ppm, no mycorrhiza were detected on the halophytes and no spores of mycorrhizal fungi were found in the soil. One percent of the roots of salt grass in soils containing 8,450 ppm of sodium were mycorrhizal. In soils containing 622 ppm of 45 percent of the roots of a salt-tolerant grass (hybrid ofAgropyron repens × Agropyron spicatum) were mycorrhizal. Halophytes such asSalicornia pacifica var.utahensis which are among the most salt tolerant halophytes of the inland salt playas rarely had mycorrhizal roots. The mycorrhizal associations appear to be very limited in inland salt playas with sodium content.     

Competition between two grass species with and without grazing over a productivity gradient

Soil nutrient-level and herbivory are predicted to have opposing effects on the allocation pattern of the competitive dominant plant species. Lower stem and higher leaf allocation are favoured when plants are grazed, whereas a higher stem allocation is favoured at high nutrient levels. Grazing by hares and geese can prevent invasion of the tall Elymus athericus, into short vegetation of Festuca rubra, at unproductive stages of salt-marsh succession but not at more productive stages. We hypothesise that the negative effect of herbivory on Elymus decreases due to increasing soil nitrogen levels and shifts the competitive balance towards this species. We tested how simulated grazing and nitrogen availability affected the competitive balance between adult plants of both grass species in a greenhouse experiment. Elymus had a higher above-ground biomass production, invested relatively more in stem and root tissue and had a larger shoot length than Festuca. The above-ground relative yield of Elymus in mixtures of both species increased with increasing nitrogen levels. This indicates that Elymus was the superior competitor at high soil fertility. Although clipping removed relatively more biomass from Elymus than from Festuca and exceeded the observed biomass removal in field conditions, it did not change the competitive balance between both species. Decreasing effects of herbivory due to increasing nitrogen levels are not a likely explanation for the invasion of Elymus in productive marshes. The results suggest that once Elymus has established it can easily invade vegetation dominated by Festuca irrespective of grazing by herbivores such as hares and geese. Herbivory by small herbivores may mainly retard the invasion of this plant by influencing establishment itself.     

Non-native competitive perennial grass impedes the spread of an invasive annual grass

Invasive plants are degrading wildlands around the globe by displacing native species, reducing biodiversity, and altering ecological functions. The current approach of applying herbicides to invasive plants in wildlands has not been effective at curtailing their expansion and, in certain circumstances, may do more harm than good. Preventing the spread of invasive species has been identified as an important strategy to protect wildlands. However, few prevention strategies have actually been tested. We hypothesized that establishing competitive vegetation next to infestations would increase the biotic resistance of the plant community to invasion and decrease the invasive species propagule pressure beyond the competitive vegetation. To evaluate this, we established twelve competitive vegetation barriers in front of invasive annual grass, Taeniatherum caput-medusae (L.) Nevski, infestations. The non-native perennial grass Agropyron desertorum (Fisch. ex Link) Schult. was seeded into plant communities adjacent to the infestations to create the competitive vegetation barriers. Soil nutrient concentrations and the spread of T. caput-medusae were compared between where A. desertorum was seeded and not seeded (control treatment) 3 years after treatment. Less T. caput-medusae and lower soil ammonium and potassium concentrations in the competitive vegetation barrier than control treatment (P ≤ 0.01) suggest that establishing competitive vegetation increased the biotic resistance of the plant communities to invasion. Taeniatherum caput-medusae cover and density in the plant communities protected by the competitive vegetation barrier (locales across the barriers from the infestations) were ~42- and 47-fold less, respectively, than unprotected plant communities (P < 0.01). This suggests that invasive plant propagule pressure was decreased in the plant communities protected by competitive vegetation barriers. The establishment of competitive vegetation around infestations may be an effective strategy to prevent or at least reduce the spread of invasive plant species.     

Soil and plant water relations in a crested wheatgrass pasture: response to spring grazing by cattle

Summary Few field studies have attempted to relate effects of actual livestock grazing on soil and plant water status. The present study was initiated to determine the effects of periodic defoliations by cattle during spring on soil moisture and plant water status in a crested wheatgrass (Agropyron cristatum (L.) Gaertn. and A. desertorum (Fisch. ex Link) Schult.) pasture in central Utah. Soil moisture in the top 130 cm of the soil profile was depleted more rapidly in ungrazed plots than in grazed plots during spring and early summer. Soil moisture depletion was more rapid in grazed plots in one paddock after 1 July due to differential regrowth, but there was no difference in soil water depletion between plots in another paddock during the same period. This difference in soil water depletion between paddocks was related to a difference in date of grazing. Although more water had been extracted from the 60 cm to 130 cm depths in ungrazed plots by late September, cumulative soil moisture depletion over the entire 193 cm profile was similar in grazed and ungrazed plots. Prior to 1 July, grazing had no effect on predawn leaf water potentials as estimated by a pressure chamber technique; however, after 1 July, predawn leaf water potentials were lower for ungrazed plants. Midday leaf water potentials were lower for grazed plants before 1 July, but did not differ between grazed and ungrazed plants after 1 July. A 4- to 8-day difference in date of defoliation did not affect either predawn or midday leaf water potentials. The observed differences in water use patterns during spring and early-summer may be important in influencing growth and competitive interactions in crested wheatgrass communities that are subject to grazing by domestic livestock.     

Establishing Native Grasses in a Big Sagebrush–Dominated Site: An Intermediate Restoration Step

Many semiarid rangelands in the Great Basin, U.S.A., are shifting dominance to woody species as a consequence of land degradation including intense livestock grazing and fire suppression. Whereas past rehabilitation efforts in Big sagebrush (Artemisia tridentata) steppes removed the shrub and added introduced forage grasses to successfully shift communities from shrublands to grasslands, current consensus is that native species should be included in restoration projects and that retention of some woody plants is desirable. We examined the potential for interseeding grasses into dense shrub communities as a precursor to thinning shrubs and releasing grasses from shrub interference. We compared seedling establishment of the native grass, Bluebunch wheatgrass (Pseudoroegneria spicata), with that of the Eurasia grass, Crested wheatgrass (Agropyron desertorum), in dense Ar. tridentata stands. Shrubs may play an important role as nurse plants for seedling establishment (reduced solar radiation, “island of fertility” effect) but result in highly contrasting light environments and root interference for seedlings. In experimental plots, we examined effects of Ar. tridentata shade levels (0, 40, 70, and 90% reduction of solar radiation) and initial root exclusion (present/absent) on the establishment and growth of P. spicata and Ag. desertorum seedlings. With this design we evaluated the interference effects of Ar. tridentata on the two grasses and identified the most beneficial microsites for grass restoration in Ar. tridentata–dominated communities. We predicted seedling survival and growth to be greater under moderate shade (40% reduction) and limited root competition than under no or strong shade conditions (0 and 90%) and unrestricted root interactions. Fifty to 85% of the P. spicata and Ag. desertorum seedlings survived the dry summer months of 1995 and 1996 and the intervening winter. Neither shading nor root exclusion from Ar. tridentata affected final seedling survival of either species. Seedling biomass of both grass species was negatively affected by initial root interactions with Ar. tridentata. However, the analysis of seedling biomass variability (coefficient of variation) indicated that in all shade and root‐exclusion treatments, some seedlings of both species developed to large individuals to survive in Ar. tridentata–dominated rangelands. Thus, the use of interseeding techniques shows promise for restoring herbaceous species in dense Ar. tridentata stands and should be given further consideration when shrub retention is an important consideration.     

Morphological Analysis of Tillering in Agropyron spicatum and Agropyron desertorum

The caespitose grasses Agropyron spicatum and Agropyron desertorumexhibit a striking difference in tillering response followingexperimental clipping treatment, with plants of A. desertorumproducing up to 18 times more tillers. The two species are similarin many aspects of their phenology and physiology. Previousexamination of current photosynthate production and levels ofstored carbohydrates indicate only slight differences betweenthe species. The possible role of three anatomical/morphologicalconstraints in controlling tillering was examined. No evidencefor such constraints was found. A basal cluster of buds is presenton the parent tillers. The mean bud number per tiller was similarfor both species and the range (3–9) was identical. Nearlyall of the bud apical meristems appeared anatomically viablethroughout the growing season and vascular development occurredto within 250 to 490 µm of the various bud apices of bothspecies. Both normal fall tillers and summer tillers producedunder clipping treatment originated from the largest, most distalbuds of the basal cluster of buds. However, precocious, morphologicallydistinctive, second-order tillers occasionally grew out fromthe smaller, most basal buds of some elongating fall tillers. Agropyron spicatum, Agropyron desertorum, bluebunch wheatgrass, crested wheatgrass, bud, tiller, tillering ability, meristematic potential, vascular development, regrowth     

The effects of shading and N status on root proliferation in nutrient patches by the perennial grass Agropyron desertorum in the field

Competition for light can affect exploitation of spatially heterogeneous soil resources. To evaluate the influence of shoot status on root growth responses in nutrient-rich soil patches, we studied the effects of shading and whole-plant nitrogen status on root growth in N-enriched and nonenriched patches by mature Agropyron desertorum plants growing in the field with below-ground competition. Roots in enriched patches had greater length to weight ratios (specific root length, SRL), indicating increased absorptive surface areas, compared with roots in control patches. Increased SRL was due to increased production and length of higher order laterals rather than morphological changes in roots of the same branching order. Although the pattern of root growth rates in patches was the same for shaded and unshaded plants, the magnitude of this response to enriched patches was damped by shading. Root relative growth rates (RGR) in N-enriched patches were reduced by more than 50% by short-term shading treatments (60% reduction in photosynthetic flux density), while root RGR in unenriched patches was unaffected by shading. Unexpectedly, plants with higher nitrogen status had greater root RGR in enriched patches than plants that had not received nitrogen supplement, again with no detectable effect on root RGR in the unenriched patches. Therefore, while both shading and plant N status affected the ability of roots to exploit enriched patches by proliferation, there was no stimulation or suppression of root growth in the unenriched, control patches. Thus, plants already under competitive pressure above ground for light and below ground for nutrients should be less able to rapidly respond to opportunities presented in nutrient patches and pulses.     

Growth and carbon allocation of Agropyron desertorum following autumn defoliation

Summary Growth and carbon allocation of a cool season tussock grass, Agropyron desertorum, following defoliation of newly initiated tillers in the autumn of 1988 and 1989 were investigated. Tiller density and mortality, reproductive shoot density, root density, biomass, individual tiller weight, carbon allocation, and soil water depletion were used to evaluate the response of A. desertorum to autumn grazing. Tiller recruitment was lower in the autumn-defoliated treatment in both years compared with the control because of the cessation of tiller development following autumn defoliation. Autumn defoliation also significantly reduced the movement of ¹³C to the roots in 1988 but not in 1989. Soils were cooler and drier in 1989. Other plant growth measurements and soil water depletion rates were not different between treatments. Autumn defoliation in 1988 did not influence tiller recruitment in the following autumn. Two consecutive years of autumn defoliation did not affect tiller overwinter mortality or peak standing crop in 1990.     

Competitive abilities of introduced and native grasses

Differencesin competitive ability may explain the maintenance of existing plantpopulationsand the invasion of new areas by plant species. We used field experiments toexamine the competitive responses of Agropyron cristatum(L.) Gaertn., an introduced C₃ grass, and Boutelouagracilis (HBK.) Lag., a native C₄ grass, and thecompetitive effects of Agropyron-dominated vegetation andsuccessional prairie. We also tested whether the outcome of competitiveinteractions varied with water availability. In each vegetation type,transplants of each species were grown under two levels of competition(presenceor absence of neighboring vegetation) and three levels of water availability(high, medium, or low). Transplant survival, growth, and biomass allocationpatterns were measured. Water availability had no effect on the measuredvariables, suggesting that both species were limited by another resource.Growthrates were affected more by competition, while survival and root: shoot ratiowere affected more by transplant species identity. In the successional prairie,neighboring vegetation suppressed the growth of Agropyrontransplants less than that of Bouteloua transplants,suggesting that Agropyron has a stronger ability to resistcompetitive suppression in that vegetation type. The spread ofAgropyron into surrounding vegetation may relate to itsability to resist competitive suppression. In theAgropyron-dominated vegetation, neighboring vegetationsuppressed the growth of both species by the same extent. However, competitionaccounted for more variation in transplant growth inAgropyron-dominated vegetation than in successionalprairie, suggesting that Agropyron has strong competitiveeffects which hinder plant growth and prevent other species from establishinginAgropyron fields.