Ecophysiological responses of two dominant grasses to altered temperature and precipitation regimes

Ecosystem responses to climate change will largely be driven by responses of the dominant species. However, if co-dominant species have traits that lead them to differential responses, then predicting how ecosystem structure and function will be altered is more challenging. We assessed differences in response to climate change factors for the two dominant C₄ grass species in tallgrass prairie, Andropogon gerardii and Sorghastrum nutans, by measuring changes in a suite of plant ecophysiological traits in response to experimentally elevated air temperatures and increased precipitation variability over two growing seasons. Maximum photosynthetic rates, stomatal conductance, water-use efficiency, chlorophyll fluorescence, and leaf water potential varied with leaf and canopy temperature as well as with volumetric soil water content (0–15 cm). Both species had similar responses to imposed changes in temperature and water availability, but when differences occurred, responses by A. gerardii were more closely linked with changes in air temperature whereas S. nutans was more sensitive to changes in water availability. Moreover, S. nutans was more responsive overall than A. gerardii to climate alterations. These results indicate both grass species are responsive to forecast changes in temperature and precipitation, but their differential sensitivity to temperature and water availability suggest that future population and community structure may vary based on the magnitude and scope of an altered climate.     

Effects of precipitation on photosynthesis and water potential in Andropogon gerardii and Schizachyrium scoparium in a southern mixed grass prairie

In grassland ecosystems, spatial and temporal variability in precipitation is a key driver of species distributions and population dynamics. We experimentally manipulated precipitation to understand the physiological basis for differences in responses of species to water availability in a southern mixed grass prairie. We focused on the performance of two dominant C₄ grasses, Andropogon gerardii Vitman and Schizachyrium scoparium (Michx.) Nash, in treatments that received ambient rainfall, half of ambient rainfall (“drought” treatment), or approximately double ambient rainfall (“irrigated” treatment). Water potentials of S. scoparium were lower than A. gerardii, suggesting superior ability to adjust to water deficit in S. scoparium. Additionally, drought reduced photosynthesis to a greater extent in A. gerardii compared to S. scoparium. Leaf-level photosynthesis rates were similar in ambient and irrigated treatments, but were significantly lower in the drought treatment. Although stomatal conductance was reduced by drought, this was not limiting for photosynthesis. Leaf δ¹³C values were decreased by drought, caused by an increase in C_i/C_a. Chlorophyll fluorescence measures indicated light-harvesting rates were highest in irrigated treatments, and were lower in ambient and drought treatments. Moreover, drought resulted in a greater proportion of absorbed photon energy being lost via thermal pathways. Reductions in photosynthesis came as a result of non-stomatal limitations in the C₄ cycle. Our results provide mechanistic support for the hypothesis that S. scoparium is more drought tolerant than A. gerardii.     

Productivity and Subordinate Species Response to Dominant Grass Species and Seed Source during Restoration

Grasses can be important regulators of species diversity and ecosystem processes in prairie systems. Although C₄ grasses are usually assumed to be ecologically similar because they are in the same functional group, there may be important differences among species or between seed sources that could impact restorations. I tested whether C₄ grass species identity, seed source, or grass species richness scales to influence aboveground net primary productivity (ANPP), resistance to weed invasion, or establishment of subordinate prairie species during restoration. Plots in western Iowa, United States, were planted with equal‐sized transplants of one of five common grass species (Panicum virgatum L., Sorghastrum nutans (L.) Nash, Andropogon gerardii Vitman, Schizachyrium scoparium (Michx.) Nash, and Bouteloua curtipendula (Michx.) Torrey) either from local seed or from cultivar seed sources. These plots were compared to plots containing all five species in mixture and to nonplanted plots. Differences in ANPP were found among species but not between cultivars and noncultivars or between monocultures and mixtures. Panicum virgatum, S. nutans, and S. scoparium were more productive than A. gerardii and B. curtipendula. Weed invasion was much higher when plots were not planted with grasses. Schizachyrium scoparium allowed greater establishment of subordinant prairie species than all other focal grass species. There were two separate mechanisms by which grasses suppressed prairie species establishment either (1) by growing tall and capturing light or (2) by quickly filling in bare space by spreading horizontally through rhizome growth in short species. These results suggest that high ANPP can be found with noncultivar plantings during the first 2 years after planting and that subordinate species establishment is most likely when shorter bunchgrasses such as S. scoparium are dominant.     

Dominant Grasses Suppress Local Diversity in Restored Tallgrass Prairie

Warm‐season (C₄) grasses commonly dominate tallgrass prairie restorations, often at the expense of subordinate grasses and forbs that contribute most to diversity in this ecosystem. To assess whether the cover and abundance of dominant grass species constrain plant diversity, we removed 0, 50, or 100% of tillers of two dominant species (Andropogon gerardii or Panicum virgatum) in a 7‐year‐old prairie restoration. Removing 100% of the most abundant species, A. gerardii, significantly increased light availability, forb productivity, forb cover, species richness, species evenness, and species diversity. Removal of a less abundant but very common species, P. virgatum, did not significantly affect resource availability or the local plant community. We observed no effect of removal treatments on critical belowground resources, including inorganic soil N or soil moisture. Species richness was inversely correlated with total grass productivity and percent grass cover and positively correlated with light availability at the soil surface. These relationships suggest that differential species richness among removal treatments resulted from treatment induced differences in aboveground resources rather than the belowground resources. Selective removal of the dominant species A. gerardii provided an opportunity for seeded forb species to become established leading to an increase in species richness and diversity. Therefore, management practices that target reductions in cover or biomass of the dominant species may enhance diversity in established and grass‐dominated mesic grassland restorations.     

Productivity responses to altered rainfall patterns in a C<Subscript>4</Subscript>-dominated grassland

Rainfall variability is a key driver of ecosystem structure and function in grasslands worldwide. Changes in rainfall patterns predicted by global climate models for the central United States are expected to cause lower and increasingly variable soil water availability, which may impact net primary production and plant species composition in native Great Plains grasslands. We experimentally altered the timing and quantity of growing season rainfall inputs by lengthening inter-rainfall dry intervals by 50%, reducing rainfall quantities by 30%, or both, compared to the ambient rainfall regime in a native tallgrass prairie ecosystem in northeastern Kansas. Over three growing seasons, increased rainfall variability caused by altered rainfall timing with no change in total rainfall quantity led to lower and more variable soil water content (0–30 cm depth), a ~10% reduction in aboveground net primary productivity (ANPP), increased root to shoot ratios, and greater canopy photon flux density at 30 cm above the soil surface. Lower total ANPP primarily resulted from reduced growth, biomass and flowering of subdominant warm-season C₄ grasses while productivity of the dominant C₄ grass Andropogon gerardii was relatively unresponsive. In general, vegetation responses to increased soil water content variability were at least equal to those caused by imposing a 30% reduction in rainfall quantity without altering the timing of rainfall inputs. Reduced ANPP most likely resulted from direct effects of soil moisture deficits on root activity, plant water status, and photosynthesis. Altered rainfall regimes are likely to be an important element of climate change scenarios in this grassland, and the nature of interactions with other climate change elements remains a significant challenge for predicting ecosystem responses to climate change.     

Growth, gas exchange, leaf nitrogen and carbohydrate concentrations in NAD‐ME and NADP‐ME C4 grasses grown in elevated CO2

Plants with the C₄ photosynthetic pathway have predominantly one of three decarboxylation enzymes in their bundle sheath cells. Within the grass family (Poaceae) bundle sheath leakiness to CO₂ is purported to be lowest in the nicotinamide adenine dinucleotide phosphate-malic enzyme (NADP-ME, EC 1.1.1.40) group, highest in the NAD-ME (EC 1.1.1.39) group and intermediate in the phosphoenolpyruvate carboxykinase (PCK, EC 4.1.1.32) group. We investigated the hypothesis that growth and photosynthesis of NAD-ME C₄ grasses would respond more to elevated CO₂ treatment than NADP-ME grasses. Plants were grown in 8-1 pots in growth chambers with ample water and fertilizer for 39 days at a continuous CO₂ concentration of either 350 or 700 µl l^?1. NAD-ME species included Bouteloua gracilis Lag. ex Steud (Blue grama), Buchloe dactyloides (Nutt.) Engelm. (Buffalo grass) and Panicum virgatum L. (Switchgrass) and the NADP-ME species were Andropogon gerardii Vittman (Big bluestem), Schizachyrium scoparium (Michx.) Nash (Little bluestem), and Sorghastrum nutans (L.) Nash (Indian grass). Contrary to our hypothesis, growth of the NADP-ME grasses was generally greater under elevated CO₂ (significant for A. gerardii and S. nutans), while none of the NAD-ME grasses had a significant growth response. Increased leaf total non-structural carbohydrate (TNC) was associated with greater growth responses of NADP-ME grasses. Decreased leaf nitrogen in NADP-ME species grown at elevated CO₂ was found to be an artifact of TNC dilution. Assimilation (A) vs intercellular CO₂ (C_i) curves revealed that leaf photosynthesis was not saturated at 350 µl l^?1 CO₂ in any of these C₄ grasses. Assimilation of elevated CO₂-grown A. gerardii was higher than in plants grown in ambient CO₂. In contrast, B. gracilis grown in elevated CO₂ displayed lower A, a trait more commonly reported in C₃ plants. Photosynthetic acclimation in B. gracilis was not related to leaf TNC or nitrogen concentrations, but A:C_i curves suggest a reduction in activity of both phosphoenolpyruvate (PEP) carboxylase (EC 4.1.1.31) and ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco, EC 4.1.1.39). Some adaptation of stomatal functioning was also seen in B. gracilis and A. gerardii leaves grown in elevated CO₂. Our study shows that C₄ grasses have the capacity for increased growth and photosynthesis under elevated CO₂ even when water and nutrients are non-limiting. While it was the NADP-ME species which had significant responses in the present study, we have previously reported significant growth increases in elevated CO₂ for B. gracilis.     

Comprehensive evaluation of the allelopathic potential of Elymus nutans

Elymus nutans has been widely planted together with other perennial grasses for rebuilding degraded alpine meadow atop the Qinghai‐Tibetan Plateau. However, the rebuilt sown pastures begin to decline a few years after establishing. One of the possible causes for the degradation of sown grassland may come from allelopathy of planted grasses. The purpose of this study was to examine allelopathic potential of Elymus nutans. Three types of aqueous extract from Elymus nutans and its root zone soil were prepared, and 5 highland crops and 5 perennial grasses were used as recipient plants. Elymus nutans exhibited strong allelopathic potential on germination and seedling growth of 5 crops, but different crops or perennial grasses respond to the extract differently. The pieces aqueous extract have stronger inhibition than whole plant extract and root zone soil extract. Hordeum vulgar var. nudum, Avena sativa, and Festuca sinensis were the most affected, while Chenopodium quinoa and Elymus sibiricus were the least affected. Elymus nutans presented less influence on Poa pratensis and Poa crymophylla than on Festuca sinensis. It is recommended that the species combination of mixture for restoration should be considered for allopathic effects on the coseeding to decrease the seeding rate ratio of Elymus nutans. The annual dicot crop seeds of Chenopodium quinoa and Brassica napus can be used as alternative subsequent crop for the seed field of Elymus nutans monoculture.     

Plant demographic responses to mycorrhizal symbiosis in tallgrass prairie

The effects of mycorrhizal symbiosis on seedling emergence, flowering and densities of several grasses and forbs were assessed in native tallgrass prairie and in sown garden populations at the Konza Prairie in northeastern Kansas. Mycorrhizal activity was experimentally suppressed with the fungicide benomyl. Flowering and stem densities of the cool-season grass, Dichanthelium oligosanthes, sedges (Carex spp.), and the forb Aster ericoides were higher in non-mycorrhizal (benomyl-treated) than in mycorrhizal plots and the magnitude of these differences was significantly affected by burning. Mycorrhizae significantly enhanced flowering of the warmseason grasses Andropogon gerardii and Sorghastrum nutans in burned prairie, but not in unburned sites. These patterns suggest that mycorrhizal effects on the dynamics of cool-season graminoid and forb populations are likely to be mediated indirectly through effects of the symbiosis on the competitive dominance of their neighbors. Seedling emergence rates of the cool-season C₃ grasses Elymus canadensis and Koeleria cristata were significantly reduced in the benomyl-treated plots, whereas benomyl treatment had no significant effect on seedling emergence of the warm-season C₄ grasses A. gerardii and Panicum virgatum. The forbs showed variable responses. Seedling emergence of Liatris aspera was greater under mycorrhizal conditions, but that of Dalea purpurea was unaffected by mycorrhizal treatment. These results show that effects of mycorrhizal symbiosis on the population dynamics of co-occurring prairie plants vary significantly both among species and among different life history stages within species. The results also indicate that mycorrhizas and fire interact to influence competitive interactions and demographic patterns of tallgrass prairie plant populations.     

Seasonal variability of forest sensitivity to heat and drought stresses: A synthesis based on carbon fluxes from North American forest ecosystems

Climate extremes such as heat waves and droughts are projected to occur more frequently with increasing temperature and an intensified hydrological cycle. It is important to understand and quantify how forest carbon fluxes respond to heat and drought stress. In this study, we developed a series of daily indices of sensitivity to heat and drought stress as indicated by air temperature (T_a) and evaporative fraction (EF). Using normalized daily carbon fluxes from the FLUXNET Network for 34 forest sites in North America, the seasonal pattern of sensitivities of net ecosystem productivity (NEP), gross ecosystem productivity (GEP) and ecosystem respiration (RE) in response to T_a and EF anomalies were compared for different forest types. The results showed that warm temperatures in spring had a positive effect on NEP in conifer forests but a negative impact in deciduous forests. GEP in conifer forests increased with higher temperature anomalies in spring but decreased in summer. The drought‐induced decrease in NEP, which mostly occurred in the deciduous forests, was mostly driven by the reduction in GEP. In conifer forests, drought had a similar dampening effect on both GEP and RE, therefore leading to a neutral NEP response. The NEP sensitivity to T_a anomalies increased with increasing mean annual temperature. Drier sites were less sensitive to drought stress in summer. Natural forests with older stand age tended to be more resilient to the climate stresses compared to managed younger forests. The results of the Classification and Regression Tree analysis showed that seasons and ecosystem productivity were the most powerful variables in explaining the variation of forest sensitivity to heat and drought stress. Our results implied that the magnitude and direction of carbon flux changes in response to climate extremes are highly dependent on the seasonal dynamics of forests and the timing of the climate extremes.     

控水与补水条件下连续热浪对闽楠光合特性和生长速率的影响

全球变暖背景下,亚热带地区极端气候热浪事件发生频率持续增加。高频热浪及其伴随的高温和干旱复合胁迫将严重影响植物的光合特性,抑制植物的生长,甚至造成死亡。然而,目前亚热带树木光合特性及生长对高频热浪及复合胁迫的响应仍不明确。以亚热带阔叶树种闽楠苗木为研究对象进行了热浪模拟实验,关注了补水和控水处理不同水分环境下连续热浪对闽楠光合特性及生长速率的影响。结果表明,补水处理下闽楠净光合速率(Pn)在单次热浪影响下显著下降了34%,同时水分利用效率显著(WUE)下降,但蒸腾速率(Tr)、气孔导度(Gs)和叶片水汽压亏缺(Leaf_vpd)显著上升(P<0.05)。表明水分充沛的高温环境中闽楠可通过增加蒸腾耗水加速水分蒸散来调节叶片的温度,增强植株光合特性对热浪的抗性。而控水处理下单次热浪处理组Tr和Gs未显著上升,以及Tr与Leaf_vpd在干湿环境下线性拟合的不同斜率,说明水分胁迫会降低叶片降温的效率,加剧热浪对闽楠光合特性的影响。水分和高温的复合胁迫还延长了闽楠光合特性在热浪后的恢复过程,高温胁迫下闽楠Pn在15 d后恢复至未干扰水平,但复合胁迫...     

Degraded soil increases the performance of a dominant grass, <Emphasis Type="Italic">Andropogon gerardii</Emphasis> (Big bluestem)

Dominant grasses can suppress subordinate species in grassland restorations. Examining factors that influence performance of a dominant grass when interacting with subordinate forbs may provide insights for maintaining plant community diversity. The objective of our study was to determine how soils of different restoration ages and functionally different forbs influence the performance (using biomass and tillering rate as proxies) of a dominant grass: Andropogon gerardii. Sites included a cultivated field and two restored prairies (4 or 16 years after restoration) at Konza Prairie (NE Kansas). We hypothesized A. gerardii performance would be greater in more degraded soils and when interacting with legumes. Soil structure, nutrient status, and microbial biomass were measured in soil that was used to conduct the plant interaction study. Andropogon gerardii performance was measured during an 18-week greenhouse experiment using the relative yield index calculated from net absolute tillering rate and final biomass measurements in three soil restoration age treatments combined with four interacting forb treatments. Restoration improved soil structure, reduced plant-available nutrients, and increased microbial biomass. Relative yield index values of A. gerardii were greater with non-legumes than legumes. Andropogon gerardii performed best in degraded soils, which may explain the difficulty in restoring tallgrass prairie diversity in long-term cultivated soil. Results from this study suggest practices that promote soil aggregation and fungal biomass, coupled with including a high abundance of legumes in seed mixes could reduce dominance of A. gerardii and likely increase plant diversity in tallgrass prairie restorations.     

Biological invasions and climate change amplify each other’s effects on dryland degradation

Climate models predict that, in the coming decades, many arid regions will experience increasingly hot conditions and will be affected more frequently by drought. These regions are also experiencing rapid vegetation change, notably invasion by exotic grasses. Invasive grasses spread rapidly into native desert ecosystems due, in particular, to interannual variability in precipitation and periodic fires. The resultant destruction of non-fire-adapted native shrub and grass communities and of the inherent soil resource heterogeneity can yield invader-dominated grasslands. Moreover, recurrent droughts are expected to cause widespread physiological stress and mortality of both invasive and native plants, as well as the loss of soil resources. However, the magnitude of these effects may differ between invasive and native grasses, especially under warmer conditions, rendering the trajectory of vegetated communities uncertain. Using the Biosphere 2 facility in the Sonoran Desert, we evaluated the viability of these hypothesized relationships by simulating combinations of drought and elevated temperature (+5°C) and assessing the ecophysiological and mortality responses of both a dominant invasive grass (Pennisetum ciliare or buffelgrass) and a dominant native grass (Heteropogan contortus or tanglehead). While both grasses survived protracted drought at ambient temperatures by inducing dormancy, drought under warmed conditions exceeded the tolerance limits of the native species, resulting in greater and more rapid mortality than exhibited by the invasive. Thus, two major drivers of global environmental change, biological invasion and climate change, can be expected to synergistically accelerate ecosystem degradation unless large-scale interventions are enacted.     

Effects of N on Plant Response to Heat-wave: A Field Study with Prairie Vegetation

More intense, more frequent, and longer heat-waves are expected in the future due to global warming, which could have dramatic ecological impacts. Increasing nitrogen (N) availability and its dynamics will likely impact plant responses to heat stress and carbon (C) sequestration in terrestrial ecosystems. This field study examined the effects of N availability on plant response to heat-stress (HS) treatment in naturally-occurring vegetation. HS (5 d at ambient or 40.5 ℃) and N treatments (±N) were applied to 16 1 m2 plots in restored prairie vegetation dominated by Andropogon gerardii (warm-season C4 grass) and Solidago canadensis (warm-season C3 forb). Before, during, and after HS, air, canopy, and soil temperature were monitored; net CO2 assimilation (Pn), quantum yield of photosystem Ⅱ (φPsⅡ), stomatal conductance (gs), and leaf water potential (Ψw) of the dominant species and soil respiration (Rsolf) of each plot were measured daily during HS. One week after HS, plots were harvested, and C% and N% were determined for rhizosphere and bulk soil, and above-ground tissue (green/senescent leaf, stem, and flower). Photosynthetic N-use efficiency (PNUE) and N resorption rate (NRR) were calculated. HS decreased Pn, gs, Ψw, and PNUE for both species, and N treatment generally increased these variables (±HS), but often slowed their poat-HS recovery. Aboveground biomass tended to decrease with HS in both species (and for green leaf mass in S. canadensis), but decrease with N for ,4. gerardii and increase with N for S. canadensis. For A. gerardii, HS tended to decrease N% in green tissues with N, whereas in S. canadensis, HS increased N% in green leaves.Added N decreased NRR for A. gerardii and HS increased NRR for S. canadensis. These results suggest that heat waves,though transient, could have significant effects on plants, communities, and ecosystem N cycling, and N can influence the effect of heat waves.     

Adaptation to high temperature mitigates the impact of water deficit during combined heat and drought stress in C3 sunflower and C4 maize varieties with contrasting drought tolerance

Heat and drought stress frequently occur together, however, their impact on plant growth and photosynthesis (P_N) is unclear. The frequency, duration and severity of heat and drought stress events are predicted to increase in the future, having severe implications for agricultural productivity and food security. To assess the impact on plant gas exchange, physiology and morphology we grew drought tolerant and sensitive varieties of C3 sunflower (Helianthus annuus) and C4 maize (Zea mays) under conditions of elevated temperature for 4 weeks prior to the imposition of water deficit. The negative impact of temperature on P_N was most apparent in sunflower. The drought tolerant sunflower retained ribulose‐1,5‐bisphosphate carboxylase/oxygenase (RubisCO) activity under heat stress to a greater extent than its drought sensitive counterpart. Maize exhibited no varietal difference in response to increased temperature. In contrast to previous studies, where a sudden rise in temperature induced an increase in stomatal conductance (G_s), we observed no change or a reduction in G_s with elevated temperature, which alongside lower leaf area mitigated the impact of drought at the higher temperature. The drought tolerant sunflower and maize varieties exhibited greater investment in root‐systems, allowing greater uptake of the available soil water. Elevated temperatures associated with heat‐waves will have profound negative impacts on crop growth in both sunflower and maize, but the deleterious effect on P_N was less apparent in the drought tolerant sunflower and both maize varieties. As C4 plants generally exhibit water use efficiency (WUE) and resistance to heat stress, selection on the basis of tolerance to heat and drought stress would be more beneficial to the yields of C3 crops cultivated in drought prone semi‐arid regions.     

Overaccumulation of glycine betaine enhances tolerance to drought and heat stress in wheat leaves in the protection of photosynthesis

We investigated the different responses of wheat (Triticum aestivum L.) plants to drought- (DS) and heat stress (HS), and analyzed the physiological mechanisms of glycine betaine (GB) involved in the improvement of wheat tolerance to the combination of these stresses. The transgenic wheat T6 line was generated by introducing a gene encoding betaine aldehyde dehydrogenase (BADH) into the wild-type (WT) Shi4185 line. The gene was cloned from the Garden Orache plant (Atriplex hortensis L.). Wheat seedlings were subjected to drought stress (30%, PEG-6000), heat stress (40°C), and their combination. Photosynthetic gas exchange, water status and lipid peroxidation of wheat leaves were examined under different stresses. When subjected to a combination of drought and heat, the inhibition of photosynthesis was significantly increased compared to that under DS or HS alone. The increased inhibition of photosynthesis by the combined stresses was not simply the additive stress effect of separate heat- and drought treatments; different responses in plant physiology to DS and HS were also found. HS decreased the chlorophyll (Chl) content, net photosynthetic rate (P _N), carboxylation efficiency (CE) and apparent quantum yield (AQY) more than DS but DS decreased the transpiration rate (E), stomata conductance (g _s) and intercellular CO₂ concentration (C _i) more than HS. GB over-accumulation led to increased photosynthesis not only under individual DS or HS but also under their combination. The enhancement of antioxidant activity and the improvement of water status may be the mechanisms underlying the improvement of photosynthesis by GB in wheat plants.     

Effects of experimental rainfall manipulations on Chihuahuan Desert grassland and shrubland plant communities

Aridland ecosystems are predicted to be responsive to both increases and decreases in precipitation. In addition, chronic droughts may contribute to encroachment of native C₃ shrubs into C₄-dominated grasslands. We conducted a long-term rainfall manipulation experiment in native grassland, shrubland and the grass–shrub ecotone in the northern Chihuahuan Desert, USA. We evaluated the effects of 5 years of experimental drought and 4 years of water addition on plant community structure and dynamics. We assessed the effects of altered rainfall regimes on the abundance of dominant species as well as on species richness and subdominant grasses, forbs and shrubs. Nonmetric multidimensional scaling and MANOVA were used to quantify changes in species composition in response to chronic addition or reduction of rainfall. We found that drought consistently and strongly decreased cover of Bouteloua eriopoda, the dominant C₄ grass in this system, whereas water addition slightly increased cover, with little variation between years. In contrast, neither chronic drought nor increased rainfall had consistent effects on the cover of Larrea tridentata, the dominant C₃ shrub. Species richness declined in shrub-dominated vegetation in response to drought whereas richness increased or was unaffected by water addition or drought in mixed- and grass-dominated vegetation. Cover of subdominant shrubs, grasses and forbs changed significantly over time, primarily in response to interannual rainfall variability more so than to our experimental rainfall treatments. Nevertheless, drought and water addition shifted the species composition of plant communities in all three vegetation types. Overall, we found that B. eriopoda responded strongly to drought and less so to irrigation, whereas L. tridentata showed limited response to either treatment. The strong decline in grass cover and the resistance of shrub cover to rainfall reduction suggest that chronic drought may be a key factor promoting shrub dominance during encroachment into desert grassland.     

Water relations and growth of three grasses during wet and drought years in a tallgrass prairie

Summary The water relations and growth of three tallgrass prairie species Panicum virgatum, Andropogon gerardii and A. scoparius were examined in irrigated and unwatered prairie in eastern Kansas (USA). Measurements of the osmotic potential at full turgor, ¹⁰⁰ , at zero turgor, ⁰, and growth of vegetative and reproductive tillers were made in a year with above-normal precipitation and a drought year to evaluate: 1) the ability of these grasses to osmotically adjust in response to water stress and 2) the effect of drought or supplemental water on growth of these species. Although these grasses adjusted osmotically even in the wet year, the degree of adjustment of ¹⁰⁰ and ⁰ in the drought year was relatively large (0.60–0.78 MPa and 0.88–1.34 MPa, respectively) compared to reports for other species. Seasonal minimum values of ¹⁰⁰ and ⁰ for these grasses in the drought year were lower than in most mesic species and seasonal fluctuations in ¹⁰⁰ and ⁰ were greater than reported for most mesic or xeric species. The relatively frequent occurrence of drought in sub-humid tallgrass prairies may partially explain the greater than expected magnitude of osmotic adjustment in these grasses.Irrigation in the wet year increased reproductive biomass in the mesic grass P. virgatum, but had no effect on A. gerardii or the more xeric grass A. scoparius. However, irrigation in the drought year increased maximum shoot biomass in all three grasses significantly with the largest increase in P. virgatum. Reproduction in P. virgatum was also increased more by irrigation in the drought year compared to the other grasses. Irrigation did not increase season's end production of A. gerardii in the wet year, but in the drought year production was 28% greater in irrigated than unwatered prairie. The combination of these water relations and growth responses of the three grasses to wetter than normal and drought years supports their reported distribution along a moisture gradient in tallgrass prairies.     

The Role of Seed and Vegetative Reproduction in Plant Recruitment and Demography in Tallgrass Prairie

Recruitment, establishment and survivorship of seed- and vegetatively-derived shoots were quantified biweekly in annually burned and infrequently burned tallgrass prairie to investigate the contributions of seed and vegetative reproduction to the maintenance and dynamics of tallgrass prairie plant populations, the demography of seedlings and ramets, and the influence of fire on the demography of grasses and forbs. Clonally produced grass and forb ramets comprised >99%of all established shoots present at the end of the growing season, whereas established seedlings accounted for <1%,emphasizing the rarity of successful seedling establishment and the importance of vegetative reproduction in driving the annual regeneration and dynamics of aboveground plant populations in tallgrass prairie. Most recruitment from vegetative reproduction occurred early in the growing season and was higher in annually burned than infrequently burned sites, although low levels of new stem recruitment occurred continuously throughout the growing season. Peak recruitment on annually burned prairie coincided with peak recruitment of the dominant C₄ grasses Andropogon gerardii and Sorghastrum nutans prior to prescribed spring fire, with a second peak in recruitment occurring following fire. On infrequently burned prairie, grass and forb recruitment was highest in early April and declined steadily through May. The naturalized C₃ grass, Poa pratensis, was responsible for most of the early recruitment on unburned sites, whereas A. gerardii contributed most to recruitment later in May. Infrequently burned prairie was dominated by these two grasses and contained a larger forb component than annually burned prairie. The principal demographic effect of fire was on ramet natality rather than mortality. Fire regime, plant functional group, or timing of cohort emergence before or after fire did not affect ramet survivorship. C₄ grass shoots that emerged early and were damaged by fire showed similar survivorship patterns to tillers that emerged after fire. Differences in species composition between annually burned and infrequently burned prairie are driven by fire effects on vegetative reproduction and appear to be related principally to the effect of fire and detritus accumulation on the development of belowground vegetative meristems of C₄ grasses and their emergence dynamics.     

Ecosystem Engineering by Plants on Wave-Exposed Intertidal Flats Is Governed by Relationships between Effect and Response Traits

In hydrodynamically stressful environments, some species—known as ecosystem engineers—are able to modify the environment for their own benefit. Little is known however, about the interaction between functional plant traits and ecosystem engineering. We studied the responses of Scirpus tabernaemontani and Scirpus maritimus to wave impact in full-scale flume experiments. Stem density and biomass were used to predict the ecosystem engineering effect of wave attenuation. Also the drag force on plants, their bending angle after wave impact and the stem biomechanical properties were quantified as both responses of stress experienced and effects on ecosystem engineering. We analyzed lignin, cellulose, and silica contents as traits likely effecting stress resistance (avoidance, tolerance). Stem density and biomass were strong predictors for wave attenuation, S. maritimus showing a higher effect than S. tabernaemontani. The drag force and drag force per wet frontal area both differed significantly between the species at shallow water depths (20 cm). At greater depths (35 cm), drag forces and bending angles were significantly higher for S. maritimus than for S. tabernaemontani. However, they do not differ in drag force per wet frontal area due to the larger plant surface of S. maritimus. Stem resistance to breaking and stem flexibility were significantly higher in S. tabernaemontani, having a higher cellulose concentration and a larger cross-section in its basal stem parts. S. maritimus had clearly more lignin and silica contents in the basal stem parts than S. tabernaemontani. We concluded that the effect of biomass seems more relevant for the engineering effect of emergent macrophytes with leaves than species morphology: S. tabernaemontani has avoiding traits with minor effects on wave attenuation; S. maritimus has tolerating traits with larger effects. This implies that ecosystem engineering effects are directly linked with traits affecting species stress resistance and responding to stress experienced.