Paul B. Hook

Plant-Soil interactions in temperate grasslands

We present a conceptual model in which plant-soil interactions in grasslands are characterized by the extent to which water is limiting. Plant-soil interactions in dry grasslands, those dominated by water limitation (‘belowground-dominance’), are fundamentally different from plant-soil interactions in subhumid grasslands, where resource limitations vary in time and space among water, nitrogen, and light (‘indeterminate dominance’). In the belowground-dominance grasslands, the strong limitation of soil water leads to complete (though uneven) occupation of the soil by roots, but insufficient resources to support continuous aboveground plant cover. Discontinuous aboveground plant cover leads to strong biological and physical forces that result in the accumulation of soil materials beneath individual plants in resource islands. The degree of accumulation in these resource islands is strongly influenced by plant functional type (lifespan, growth form, root:shoot ratio, photosynthetic pathway), with the largest resource islands accumulating under perennial bunchgrasses. Resource islands develop over decadal time scales, but may be reduced to the level of bare ground following death of an individual plant in as little as 3 years. These resource islands may have a great deal of significance as an index of recovery from disturbance, an indicator of ecosystem stability or harbinger of desertification, or may be significant because of possible feedbacks to plant establishment. In the grasslands in which the dominant resource limiting plant community dynamics is indeterminate, plant cover is relatively continuous, and thus the major force in plant-soil interactions is related to the feedbacks among plant biomass production, litter quality and nutrient availability. With increasing precipitation, the over-riding importance of water as a limiting factor diminishes, and four other factors become important in determining plant community and ecosystem dynamics: soil nitrogen, herbivory, fire, and light. Thus, several different strategies for competing for resources are present in this portion of the gradient. These strategies are represented by different plant traits, for example root:shoot allocation, height and photosynthetic pathway type (C3 vs. C4) and nitrogen fixation, each of which has a different influence on litter quality and thus nutrient availability. Recent work has indicated that there are strong feedbacks between plant community structure, diversity, and soil attributes including nitrogen availability and carbon storage. Across both types of grasslands, there is strong evidence that human forces that alter plant community structure, such as invasions by nonnative annual plants or changes in grazing or fire regime, alters the pattern, quantity, and quality of soil organic matter in grassland ecosystems. The reverse influence of soils on plant communities is also strong; in turn, alterations of soil nutrient supply in grasslands can have major influences on plant species composition, plant diversity, and primary productivity.

BIOGEOCHEMISTRY IN A SHORTGRASS LANDSCAPE: CONTROL BY TOPOGRAPHY, SOIL TEXTURE, AND MICROCLIMATE

Biogeochemistry of terrestrial ecosystems is controlled by interactions among factors operating at several spatial and temporal scales. The purpose of this study was to evaluate the relative importance and interaction of relatively static landscape factors and more dynamic factors in a shortgrass steppe landscape. The landscape factors examined were topographic position, and soil texture. The dynamic factors studied were seasonal climate and the localized effects of individual plants on soils. Patterns were evaluated by sampling soil between and under individual Bouteloua gracilis plants in paired upland (erosional) and lowland (depositional) plots at eight locations at the Central Plains Experimental Range (CPER), Colorado. We quantified five organic C and N pools (total, fine and coarse particulate organic matter [POM], mineral-associated organic matter [MAOM], and potentially mineralizable C and N), and we estimated seasonal patterns of in situ N dynamics with three methods (extractable inorganic N, ne...

Spatial patterns of roots in a semiarid grassland: Abundance of canopy openings and regeneration gaps

1 Spatial patterns of root biomass and plant cover were quantified in 10 late-successional, shortgrass steppe communities in which a large proportion of the soil is bare and regeneration is frequently limited by soil water. Our main objectives were to evaluate how patterns of root density were associated with previously documented variation in recruitment in canopy openings of different sizes and to estimate the abundance of openings with low root density. 2 Root biomass in the top 30 cm of soil was much lower in openings of all sizes than under plants and declined steeply as opening size increased. Biomass of light-coloured roots (i.e. those presumed to be functional) in centres of 10-, 20and 60-cm openings was 62, 33 and 4%, respectively, of that under plants. 3 Openings more than 5 cm across made up 34% of the surface. Most were small: 86% of openings were 50 cm across), which are known to enhance regeneration, had low root density. Such openings (2% of total) occupied 2% of the area and nearly all were caused by disturbance. However, many openings caused by disturbance were 30-50 cm across, a size range where interference from neighbours changes from strong to weak, and some were even smaller. Most of the area (>99.5%) was within the range of the root system of the dominant grass, Bouteloua gracilis. 5 We infer that most openings large enough to support enhanced recruitment are explored by roots of dominant bunchgrasses and that gap dynamics in shortgrass steppe involves constraints on water use in B. gracilis root systems. Because large openings are rare, variation in below-ground competition in the abundant, smaller openings may be important for regeneration.

Effects of the Invasive Forb Centaurea maculosa on grassland Carbon and Nitrogen Pools in Montana, USA

Invasions by exotic forbs are changing large areas of North American grasslands, but their biogeochemical impacts are not well characterized. Additionally, although many invasive plants may alter biogeochemistry, an invasive species’ effects have rarely been evaluated across physically diverse sites. We sampled nine sites containing the perennial Eurasian forb Centaurea maculosa to determine if this invasive species alters soil C and N pools in native grasslands in Montana, USA. We sampled surface soil in adjacent microsites with C. maculosa and native grasses and analyzed soil C and N pools with slow to rapid turnover. None of the pools evaluated in the laboratory showed significant differences between C. maculosa and grass microsites when analyzed across all sites. Some differences were found at individual sites, but they were infrequent and inconsistent: Four sites had no differences, four had differences in one or two pools with intermediate (particulate organic matter C or N) or rapid turnover rates (potentially mineralizable N), and just one site had differences encompassing pools with rapid, intermediate, and slow (total C and N, silt-and-clay-associated N) turnover rates. Where they differed, pools were usually smaller under C. maculosa plants than under native grasses, but the opposite was found at one site. In situ N availability, estimated using ion exchange resins, was significantly lower under C. maculosa than under grasses at one of three sites sampled. Results indicate that C. maculosa may sometimes reduce soil C and N pools, including those related to N availability, but they argue against generalizing about the impacts of C. maculosa in grasslands.

Ammonium Removal in Constructed Wetland Microcosms as Influenced by Season and Organic Carbon Load

Abstract We evaluated ammonium nitrogen removal and nitrogen transformations in three-year-old, batch-operated, subsurface wetland microcosms. Treatments included replicates of Typha latifolia, Carex rostrata, and unplanted controls when influent carbon was excluded, and C. rostrata with an influent containing organic carbon. A series of 10-day batch incubations were conducted over a simulated yearlong cycle of seasons. The presence of plants significantly enhanced ammonium removal during both summer (24°C, active plant growth) and winter (4°C, plant dormancy) conditions, but significant differences between plant species were evident only in summer when C. rostrata outperformed T. latifolia. The effect of organic carbon load was distinctly seasonal, enhancing C. rostrata ammonium removal in winter but having an inhibitory effect in summer. Season did not influence ammonium removal in T. latifolia or unplanted columns. Net production of organic carbon was evident year-round in units without an influent organic carbon source, but was enhanced in summer, especially for C. rostrata, which produced significantly more than T. latifolia and unplanted controls. No differences in production were evident between species in winter. COD values for C. rostrata microcosms with and without influent organic carbon converged within 24 hours in winter and 7 days in summer. Gravel sorption, microbial immobilization and sequential nitrification/denitrification appear to be the major nitrogen removal mechanisms. All evidence suggests differences between season and species are due to differences in seasonal variation of root-zone oxidation.

Temperature, plant species and residence time effects on nitrogen removal in model treatment wetlands

Total nitrogen (TN) removal in treatment wetlands (TWs) is challenging due to nitrogen cycle complexity and the variation of influent nitrogen species. Plant species, season, temperature and hydraulic loading most likely influence root zone oxygenation and appurtenant nitrogen removal, especially for ammonium-rich wastewater. Nitrogen data were collected from two experiments utilizing batch-loaded (3-, 6-, 9- and 20-day residence times), sub-surface TWs monitored for at least one year during which temperature was varied between 4 and 24 °C. Synthetic wastewater containing 17 mg/l N as NH4 and 27 mg/l amino-N, 450 mg/l chemical oxygen demand (COD), and 13 mg/l SO4-S was applied to four replicates of Carex utriculata, Schoenoplectus acutus and Typha latifolia and unplanted controls. Plant presence and species had a greater effect on TN removal than temperature or residence time. Planted columns achieved approximately twice the nitrogen removal of unplanted controls (40-95% versus 20-50% removal) regardless of season and temperature. TWs planted with Carex outperformed both Typha and Schoenoplectus and demonstrated less temperature dependency. TN removal with Carex was excellent at all temperatures and residence times; Schoenoplectus and Typha TN removal improved at longer residence times. Reductions in TN were not accompanied by increases in NO3, which was consistently below 1 mg/l N.