Elevated CO2 further lengthens growing season under warming conditions.

Observations of a longer growing season through earlier plant growth in temperate to polar regions have been thought to be a response to climate warming. However, data from experimental warming studies indicate that many species that initiate leaf growth and flowering earlier also reach seed maturation and senesce earlier, shortening their active and reproductive periods. A conceptual model to explain this apparent contradiction, and an analysis of the effect of elevated CO2--which can delay annual life cycle events--on changing season length, have not been tested. Here we show that experimental warming in a temperate grassland led to a longer growing season through earlier leaf emergence by the first species to leaf, often a grass, and constant or delayed senescence by other species that were the last to senesce, supporting the conceptual model. Elevated CO2 further extended growing, but not reproductive, season length in the warmed grassland by conserving water, which enabled most species to remain active longer. Our results suggest that a longer growing season, especially in years or biomes where water is a limiting factor, is not due to warming alone, but also to higher atmospheric CO2 concentrations that extend the active period of plant annual life cycles.

Carbon and water relations of Salix monticola in response to winter browsing and changes in surface water hydrology: an isotopic study using δ13C and δ18O.

To ascertain whether browsing or hydrologic conditions influence the physiological performance of Salix and whether Salix and graminoids (Carex) use and possibly compete for similar water resources, we quantified the in situ seasonal patterns of plant water and carbon relations over three growing seasons. Our studies were designed to address the physiological factors which may be responsible for poor woody plant regeneration in montane riparian habitats of Rocky Mountain National Park, Colo. As these systems act to insure the delivery of fresh water to downstream users, the maintenance of their integrity is critical. We quantified plant water potentials, instantaneous rates of carbon fixation, leaf carbon isotope discrimination (Δ), leaf nitrogen content and water sources using stable isotopes of water (δ18O). The carbon and water relations of Salix were significantly affected by winter browsing by elk and in some cases by landscape position with regard to proximity to active streams. Winter browsing of Salix by elk significantly increased summer plant water potentials and integrative measures of gas exchange (Δ), though browsing did not consistently affect instantaneous rates of photosynthesis, leaf nitrogen or the sources of water used by Salix. No effect of experimental manipulations of surface water conditions on Salix physiology was observed, likely due to the mesic nature of our study period. Using a two-member linear mixing model, from δ18O values we calculated that Salix appears to rely on streams for approximately 80% of its water. In contrast, the graminoid Carex derives almost 50% of its water from rainfall, indicating divergent water source use by these two life forms. Based on these findings, winter browsing by elk improved Salix water balance possibly by altering the shoot to root ratio which in turn leads to higher water potentials and higher degrees of season-long gas exchange, while experimental damming had in general no effect on the physiological performance of Salix plants. In addition, as the water sources of Salix and Carex were significantly different, competition for water may not influence the growth, development, and regeneration of Salix. Thus, under the conditions of our study, herbivory had a positive effect on the physiological performance of Salix, but it is still unclear whether these changes in physiology transcend into improved Salix regeneration and survivorship. However, under drier environmental conditions such as lower snowpacks and lower stream flows, the browsing resistance of Salix and ecosystem regeneration may be greatly hindered because the reliance of Salix on stream water makes it vulnerable to changes in surface water and hydrological conditions.

Grazing and Plant Performance.

Grazing is more than just defoliation of plants. The impact of herbivory affects ecosystem structure and function, both above and below ground. Ultimately, effects of herbivory are expressed to varying degrees at many levels of the ecosystem. Herbivory has been shown to affect plant physiology, morphology, and genetics. Plants have evolved many ways to avoid or tolerate herbivory. Whether plant overcompensate, equally compensate, or undercompensate to herbivory depends on pre- and post-harvest conditions of the plants and their environment. To be important to the manager, the magnitude of compensation must be greater than the inherent "noise" in the system. Natural resources managers use scientific information about herbivory to reduce ambiguity in decision-making in an environment of uncertainty. If an ecological response like compensation is to have practical application for the manager, the meaningful effects must occur on time and spatial scales that the manager can respond to with available resources.

Water status of soil and vegetation in a shortgrass steppe.

In an attempt to describe some major relationships between soil and plant compartments in a shortgrass steppe, the process of water loss from the system and plant water relations throughout a drying cycle were studied. The water supply was manipulated and some soil and plant variables monitored throughout a drying cycle. Leaf conductance and leaf water potential of blue grama (Bouteloua gracilis) were measured periodically at predawn and noon. Soil water content and water potential of different layers were also monitored.Three different periods were distinguished in the water loss process throughout a drying cycle. These distinctions were made taking into account the relative contribution of different soil layers. Leaf conductance and water potential at noon slowly declined throughout the first 50 days of plant growth. After that, they rapidly decreased, reaching values of 0.29 mm s-1 and-5.0 MPa, respectively. The predawn leaf water potential remained unchanged around-0.5 MPa during the first 45 days, then rapidly decreased. This occurred when soil water of the wettest soil layer was near depletion.Predawn leaf water potentials were highly correlated with water potentials of the wettest layer. Leaf conductance and water potential at noon were correlated with effective soil water potential (soil water potential weighted by the root distribution in the profile). We concluded that root surface area limited the water flow through an important part of the day in this semiarid ecosystem. Axial root resistance did not appear important in determining the equilibrium status between leaves and the wettest soil layer.

14C distribution and utilization in blue grama as affected by temperature, water potential and defoliation regimes.

Water stress and temperature effects on growth, translocation and reallocation of 14C assimilated by blue grama (Bouteloua gracilis (H.B.K.) Lag. ex Steud.) were studied for sods extracted from shortgrass prairie. The sods were kept at either 24°/16° or 34°/16°C (day/nigh) temperatures and were labeled at two phenological stages. Three soil water potential (Ψ s) regimes of approximately 0 to -1, 0 to -15, and 0 to -30 bars were maintained by wetting and drying cycles. Reproductive plants retained more 14C in aboveground organs, used more assimilated 14C for respiration, and allocated a higher proportion of photosynthate to the labile fraction than did plants labeled at a vegetative stage. Low temperature and a Ψ s of 0 to -30 bars resulted in greater 14C translocation to belowground organs, but a larger proportion of the 14C went into structural components. Sods stressed to-30 bars and maintained at higher temperatures had higher respiration losses of 14C. No significant differences in allocation and respiration uses of 14C were found among sods grown at field capacity and at 0 to-15 bars Ψ s.During regrowth of clipped sods, more than 60% of the labile 14C in belowground organs was respired within four weeks. Higher respiration losses of labile 14C were found in severely water stressed sods that regrew under the higher temperature regime. Use of labile 14C for regrowth of new foliage was greatest for sods kept under conditions of high temperatures and optimum to moderate Ψ s. Height growth and biomass increas of new foliage were significantly less for sods stresses to -30 bars. No significant differences in reallocation or respiration losses of 14C, growth, and biomass increases were observed for sods maintained at field capacity and 0 to -15 bars. These results suggested that physiological processes in blue grama were affected and could not recover fully when plants were subjected to severe water stress. However, upon relief of stress, those sods maintained at a moderate soil water stress level to 0 to -15 bars were capable of rapid recovery. Significance of these findings is discussed in the context of evolutionary success of blue grama in a semi-arid environment.