Accelerated development in Johnsongrass seedlings (Sorghum halepense) suppresses the growth of native grasses through size-asymmetric competition.

Invasive plant species often dominate native species in competition, augmenting other potential advantages such as release from natural enemies. Resource pre-emption may be a particularly important mechanism for establishing dominance over competitors of the same functional type. We hypothesized that competitive success of an exotic grass against native grasses is mediated by establishing an early size advantage. We tested this prediction among four perennial C4 warm-season grasses: the exotic weed Johnsongrass (Sorghum halepense), big bluestem (Andropogon gerardii), little bluestem (Schizachyrium scoparius) and switchgrass (Panicum virgatum). We predicted that a) the competitive effect of Johnsongrass on target species would be proportional to their initial biomass difference, b) competitive effect and response would be negatively correlated and c) soil fertility would have little effect on competitive relationships. In a greenhouse, plants of the four species were grown from seed either alone or with one Johnsongrass neighbor at two fertilizer levels and periodically harvested. The first two hypotheses were supported: The seedling biomass of single plants at first harvest (50 days after seeding) ranked the same way as the competitive effect of Johnsongrass on target species: Johnsongrass < big bluestem < little bluestem/switchgrass, while Johnsongrass responded more strongly to competition from Johnsongrass than from native species. At final harvest, native plants growing with Johnsongrass attained between 2-5% of their single-plant non-root biomass, while Johnsongrass growing with native species attained 89% of single-plant non-root biomass. Fertilization enhanced Johnsongrass' competitive effects on native species, but added little to the already severe competitive suppression. Accelerated early growth of Johnsongrass seedlings relative to native seedlings appeared to enable subsequent resource pre-emption. Size-asymmetric competition and resource-pre-emption may be a critical mechanism by which exotic invasive species displace functionally similar native species and alter the functional dynamics of native communities.

[Challenges and probable solutions for using stable isotope techniques to identify plant water sources in karst regions: A review]. / é‡‡ç”¨ç¨³å®šåŒä½ç´ æŠ€æœ¯åˆ¤å®šå–€æ–¯ç‰¹åœ°åŒºæ¤ç‰©æ°´åˆ†æ¥æºçš„æŒ‘æˆ˜ä¸Žå¯èƒ½åº”å¯¹æ–¹æ¡ˆ.

Karst regions, which account for about 15% of the terrestrial surface area, are characte-rized by specific hydrogeological structure different from most non-karst regions. Thus, many research methods that are used in non-karst regions cannot be directly used in karst regions. This issue is especially relevant to research on plant water sources. In this paper, origins and possible solutions to the common problems associated with research on water sources used by karst plant species were reviewed. Four questions were addressed: 1) why is it important to determine plant water source in karst regions? 2) Why are stable isotopes used? 3) What are the challenges associated with using stable isotopes in karst regions? 4) What are the probable solutions for these challenges? This review emphasized the advantages of using stable isotope techniques to identify sources of water used by karst plant species and the challenges associated with satisfying the prerequisites of this method. It is suggested that sources of water used by plant species in karst regions need not to be divided into specific depths and the method of identifying sources of water used by plant species based on their hydrologic properties was much applicable.

Hydraulic responses to extreme drought conditions in three co-dominant tree species in shallow soil over bedrock.

An important component of the hydrological niche involves the partitioning of water sources, but in landscapes characterized by shallow soils over fractured bedrock, root growth is highly constrained. We conducted a study to determine how physical constraints in the root zone affected the water use of three tree species that commonly coexist on the Edwards Plateau of central Texas; cedar elm (Ulmus crassifolia), live oak (Quercus fusiformis), and Ashe juniper (Juniperus ashei). The year of the study was unusually dry; minimum predawn water potentials measured in August were -8 MPa in juniper, less than -8 MPa in elm, and -5 MPa in oak. All year long, species used nearly identical water sources, based on stable isotope analysis of stem water. Sap flow velocities began to decline simultaneously in May, but the rate of decline was fastest for oak and slowest for juniper. Thus, species partitioned water by time when they could not partition water by source. Juniper lost 15-30 % of its stem hydraulic conductivity, while percent loss for oak was 70-75 %, and 90 % for elm. There was no tree mortality in the year of the study, but 2 years later, after an even more severe drought in 2011, we recorded 34, 14, 6, and 1 % mortality among oak, elm, juniper, and Texas persimmon (Diospyros texana), respectively. Among the study species, mortality rates ranked in the same order as the rate of sap flow decline in 2009. Among the angiosperms, mortality rates correlated with wood density, lending further support to the hypothesis that species with more cavitation-resistant xylem are more susceptible to catastrophic hydraulic failure under acute drought.

The water relations of two evergreen tree species in a karst savanna.

The ecohydrology of karst has not received much attention, despite the disproportionally large contribution of karst aquifers to freshwater supplies. Karst savannas, like many savannas elsewhere, are encroached by woody plants, with possibly negative consequences on aquifer recharge. However, the role of savanna tree species in hydrological processes remains unclear, not least because the location and water absorption zones of tree roots in the spatially complex subsurface strata are unknown. This study examined the water sources and water relations of two savanna trees, Quercus fusiformis (Small) and Juniperus ashei (Buchholz) in the karst region of the eastern Edwards Plateau, Texas (USA). Stable isotope analysis of stem water revealed that both species took up evaporatively enriched water during the warm season, suggesting a relatively shallow water source in the epikarst, the transition zone between soil and bedrock. Q. fusiformis had consistently higher predawn water potentials than J. ashei during drought, and thus was probably deeper-rooted and less capable of maintaining gas exchange at low water potentials. Although the water potential of both species recovered after drought-breaking spring and summer rain events, associated shifts in stem water isotope ratios did not indicate significant uptake of rainwater from the shallow soil. A hypothesis is developed to explain this phenomenon invoking a piston-flow mechanism that pushes water stored in macropores into the active root zones of the trees. Epikarst structure varied greatly with parent material and topography, and had strong effects on seasonal fluctuations in plant water status. The study suggests that tree species of the Edwards Plateau do not commonly reduce aquifer recharge by tapping directly into perched water tables, but more likely by reducing water storage in the epikarst. A more general conclusion is that models of savanna water relations based on Walter's two-layer model may not apply unequivocally to karst savannas.

Hierarchy of responses to resource pulses in arid and semi-arid ecosystems.

In arid/semi-arid ecosystems, biological resources, such as water, soil nutrients, and plant biomass, typically go through periods of high and low abundance. Short periods of high resource abundance are usually triggered by rainfall events, which, despite of the overall scarcity of rain, can saturate the resource demand of some biological processes for a time. This review develops the idea that there exists a hierarchy of soil moisture pulse events with a corresponding hierarchy of ecological responses, such that small pulses only trigger a small number of relatively minor ecological events, and larger pulses trigger a more inclusive set and some larger ecological events. This framework hinges on the observation that many biological state changes, where organisms transition from a state of lower to higher physiological activity, require a minimal triggering event size. Response thresholds are often determined by the ability of organisms to utilize soil moisture pulses of different infiltration depth or duration. For example, brief, shallow pulses can only affect surface dwelling organisms with fast response times and high tolerance for low resource levels, such as some species of the soil micro-fauna and -flora, while it takes more water and deeper infiltration to affect the physiology, growth or reproduction of higher plants. This review first discusses how precipitation, climate and site factors translate into soil moisture pulses of varying magnitude and duration. Next, the idea of the response hierarchy for ecosystem processes is developed, followed by an exploration of the possible evolutionary background for the existence of response thresholds to resource pulses. The review concludes with an outlook on global change: does the hierarchical view of precipitation effects in ecosystems provide new perspectives on the future of arid/semiarid lands?

Dominant cold desert plants do not partition warm season precipitation by event size.

We conducted experiments to examine the quantitative relationships between rainfall event size and rainwater uptake and use by four common native plant species of the Colorado Plateau, including two perennial grasses, Hilaria jamesii (C(4)) and Oryzopsis hymenoides (C(3)), and two shrubs, Ceratoides lanata (C(3)), and Gutierrezia sarothrae (C(3)). Specifically, we tested the hypothesis that grasses use small rainfall events more efficiently than shrubs and lose this advantage when events are large. Rainfall events between 2 and 20 mm were simulated in spring and summer by applying pulses of deuterium-labeled irrigation water. Afterwards, pulse water fractions in stems and the rates of leaf gas exchange were monitored for 9 days. Cumulative pulse water uptake over this interval (estimated by integrating the product of pulse fraction in stem water and daytime transpiration rate over time) was approximately linearly related to the amount of pulse water added to the ground in all four species. Across species, consistently more pulse water was taken up in summer than in spring. Relative to their leaf areas, the two grass species took up more pulse water than the two shrub species, across all event sizes and in both seasons, thus refuting the initial hypothesis. In spring, pulse water uptake did not significantly increase photosynthetic rates and in summer, pulse water uptake had similar, but relatively small effects on the photosynthetic rates of the three C(3) plants, and a larger effect on the C(4) plant H. jamesii. Based on these data, we introduce an alternative hypothesis for the responses of plant functional types to rainfall events of different sizes, building on cost-benefit considerations for active physiological responses to sudden, unpredictable changes in water availability.

Deuterium enriched irrigation indicates different forms of rain use in shrub/grass species of the Colorado Plateau.

We contrasted the seasonal use of simulated large rain events (24 mm) by three native species of the arid Colorado Plateau: the perennial grass Hilaria jamesii and two shrubs Artemesia filifolia and Coleogyne ramosissima. Deuterium-enriched water was used to distinguish shallow "pulse" water from water in deeper soil layers that were unaffected by the water input. We also measured the leaf gas exchange rates of watered and unwatered control plants for 5 days after the rain event. H. jamesii had twice the pulse water proportion in its xylem than the two shrubs in spring (approx. 70% vs 35%). In summer, the pulse water proportions of all species were around 70%. The increase in the relative pulse water uptake of the two shrubs was caused primarily by a reduction in the rate of water uptake from deeper sources, consistent with the decrease in the availability of stored winter water. Rain increased the rates of gas exchange in C. ramosissima in both seasons, in H. jamesii only in summer and had no significant effect on A. filifolia. In H. jamesii, summer rain also increased water use efficiency. This suggests three principle mechanisms for rainwater use: (1) immediate increase in gas exchange via stomatal opening (C. ramisissima), (2) immediate increase in water use efficiency through restoration of the photosynthetic apparatus (H. jamesii) and (3) conservation of deeper soil water, potentially extending photosynthetic activity into later drought periods (A. filifolia). On a ground-area basis, A. filifolia was by far the largest consumer of spring and summer rain, due to its greater ground cover, while rain use by H. jamesii was negligible. We hypothesize that a population's fraction of the total community Leaf Area Index, more than species identity, determines which species takes up most of the spring and summer precipitation and we discuss this idea in the context of Walter and Stadelmann's (1974, In: Brown JW Jr (ed) Desert biology. Academic Press, New York, pp 213-310) water partitioning hypothesis.

Mechanisms determining the degree of size asymmetry in competition among plants.

When plants are competing, larger individuals often obtain a disproportionate share of the contested resources and suppress the growth of their smaller neighbors, a phenomenon called size-asymmetric competition. We review what is known about the mechanisms that give rise to and modify the degree of size asymmetry in competition among plants, and attempt to clarify some of the confusion in the literature on size asymmetry. We broadly distinguish between mechanisms determined primarily by characteristics of contested resource from those that are influenced by the growth and behavior of the plants themselves. To generate size asymmetric resource competition, a resource must be "pre-emptable." Because of its directionality, light is the primary, but perhaps not the only, example of a pre-emptable resource. The available data suggest that competition for mineral nutrients is often size symmetric (i.e., contested resources are divided in proportion to competitor sizes), but the potential role of patchily and/or episodically supplied nutrients in causing size asymmetry is largely unexplored. Virtually nothing is known about the size symmetry of competition for water. Plasticity in morphology and physiology acts to reduce the degree of size asymmetry in competition. We argue that an allometric perspective on growth, allocation, resource uptake, and resource utilization can help us understand and quantify the mechanisms through which plants compete.