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Commissioned Review. Carbon: freshwater plants

δ13C values for freshwater aquatic plant matter varies from −11 to −50‰ and is not a clear indicator of photosynthetic pathway as in terrestrial plants. Several factors affect δ13C of aquatic plant matter. These include: (1) The δ13C signature of the source carbon has been observed to range from +1‰ for HCO3− derived from limestone to −30‰ for CO2 derived from respiration. (2) Some plants assimilate HCO3−, which is –7 to –11‰ less negative than CO2. (3) C3, C4, and CAM photosynthetic pathways are present in aquatic plants. (4) Diffusional resistances are orders of magnitude greater in the aquatic environment than in the aerial environment. The greater viscosity of water acts to reduce mixing of the carbon pool in the boundary layer with that of the bulk solution. In effect, many aquatic plants draw from a finite carbon pool, and as in terrestrial plants growing in a closed system, biochemical discrimination is reduced. In standing water, this factor results in most aquatic plants having a δ13C value similar to the source carbon. Using Farquhar's equation and other physiological data, it is possible to use δ13C values to evaluate various parameters affecting photosynthesis, such as limitations imposed by CO2 diffusion and carbon source.

Plant, Cell & Environment

Variation in plant defense suppresses herbivore performance

Defensive variability of crops and natural systems can alter herbivore communities and reduce herbivory . However, it is still unknown how defense variability translates into herbivore suppression. Nonlinear averaging and constraints in physiological tracking (also more generally called time-dependent effects) are the two mechanisms by which defense variability might impact herbivores . We conducted a set of experiments manipulating the mean and variability of a plant defense, showing that defense variability does suppress herbivore performance and that it does so through physiological tracking effects that cannot be explained by nonlinear averaging. While nonlinear averaging predicted higher or the same herbivore performance on a variable defense than on an invariable defense, we show that variability actually decreased herbivore performance and population growth rate. Defense variability reduces herbivore performance in a way that is more than the average of its parts. This is consistent with constraints in physiological matching of detoxification systems for herbivores experiencing variable toxin levels in their diet and represents a more generalizable way of understanding the impacts of variability on herbivory . Increasing defense variability in croplands at a scale encountered by individual herbivores can suppress herbivory, even if that is not anticipated by nonlinear averaging.

Current Biology

Effects of boron on growth and physiology in mallard ducklings

High concentrations of boron (B) have been associated with irrigation drainwater and aquatic plants consumed by waterfowl. Day-old mallard ( Anas platyrhynchos ) ducklings received an untreated diet (controls) or diets containing 100, 400 or 1,600 ppm B as boric acid. Survival, growth and food consumption were measured for 10 weeks. At termination, blood and tissue samples were collected for biochemical assays and histological examination. The highest dietary concentration of B caused 10% mortality, decreased overall growth and the rate of growth (sexes combined), whereas lower concentrations of B altered growth only in females. Food consumption was lower during the first 3 weeks in the 1,600-ppm group and during the second week in all B-treated groups compared to controls. Hematocrit and hemoglobin were lower and plasma calcium concentration higher in the 1,600-ppm group compared to controls. Plasma triglyceride concentration was elevated in all B-treated groups. Brain B concentration increased to 25 times that of controls in the 1,600-ppm group. Brain ATP decreased with increasing dietary B. Brain acetylcholinesterase activity and total ATPase activity (in males) were elevated and protein concentration lowered in the 1,600-ppm group. Boron accumulated less in the liver than in the brain but resulted in an initial elevation of hepatic glutathione. These findings, in combination with altered duckling behavior, suggest that concentrations of B occurring in aquatic plants could adversely affect normal duckling development.

Environmental Toxicology and Chemistry

Ammonia and aquatic ecosystems – A review of global sources, biogeochemical cycling, and effects on fish

The purpose of this review is to better understand the full life cycle and influence of ammonia from an aquatic biology perspective. While ammonia has toxic properties in water and air, it also plays a central role in the biogeochemical nitrogen (N) cycle and regulates mechanisms of normal and abnormal fish physiology. Additionally, as the second most synthesized chemical on Earth, ammonia contributes economic value to many sectors, particularly fertilizers, energy storage, explosives, refrigerants, and plastics. But, with so many uses, industrial N 2 -fixation effectively doubles natural reactive N concentrations in the environment. The consequence is global, with excess fixed nitrogen driving degradation of soils, water, and air; intensifying eutrophication, biodiversity loss, and climate change; and creating health risks for humans, wildlife, and fisheries. Thus, the need for ammonia research in aquatic systems is growing. In response, we prepared this review to better understand the complexities and connectedness of environmental ammonia. Even the term “ammonia” has multiple meanings. So, we have clarified the nomenclature, identified units of measurement, and summarized methods to measure ammonia in water. We then discuss ammonia in the context of the N-cycle, review its role in fish physiology and mechanisms of toxicity, and integrate the effects of human N-fixation, which continuously expands ammonia's sources and uses. Ammonia is being developed as a carbon-free energy carrier with potential to increase reactive nitrogen in the environment. With this in mind, we review the global impacts of excess reactive nitrogen and consider the current monitoring and regulatory frameworks for ammonia. The presented synthesis illustrates the complex and interactive dynamics of ammonia as a plant nutrient, energy molecule, feedstock, waste product, contaminant, N-cycle participant, regulator of animal physiology, toxicant, and agent of environmental change. Few molecules are as influential as ammonia in the management and resilience of Earth's resources.

Science of the Total Environment

Effects of arsenate on growth and physiology in mallard ducklings

Arsenic (As) has been found at elevated concentrations in irrigation drainwater and in aquatic plants utilized by waterfowl. Mallard ( Anas platyrhynchos ) ducklings received an untreated diet (controls) or diets containing 30, 100 or 300 ppm As added as sodium arsenate. After 10 weeks blood and tissue samples were collected for biochemical and histological examination. Arsenic accumulated significantly in brain and liver of ducklings fed 100 or 300 ppm but did not result in histopathological lesions. The 300-ppm dietary As concentration decreased overall growth (weight gain) in males, whereas all concentrations of As decreased overall growth and rate of growth in females. Food consumption was less during the first three weeks in the 300-ppm group and during the second week for the 100-ppm group compared to controls. Plasma sorbitol dehydrogenase activity and plasma glucose concentration were higher in the 300-ppm group compared to controls. Plasma triglyceride concentration increased in all As-treated groups. Brain ATP was lower in the 300-ppm group and sodium/potassium-dependent ATPase activity was higher in the 30- and 100-ppm groups. Hepatic glutathione peroxidase activity was lower in the 300-ppm group and malondialdehyde lower in all treatment groups. All treatment levels caused elevation in hepatic glutathione and ATP concentrations. These findings, in combination with altered duckling behavior (increased resting time), suggest that concentrations of As that have been found in aquatic plants (up to 430 ppm dry weight) could adversely affect normal duckling development.

Environmental Toxicology and Chemistry

Spatial and temporal trends of drought effects in a heterogeneous semi-arid forest ecosystem

Drought has long been recognized as a driving mechanism in the forests of western North America and drought-induced mortality has been documented across genera in recent years. Given the frequency of these events are expected to increase in the future, understanding patterns of mortality and plant response to severe drought is important to resource managers. Drought can affect the functional, physiological, structural, and demographic properties of forest ecosystems. Remote sensing studies have documented changes in forest properties due to direct and indirect effects of drought; however, few studies have addressed this at local scales needed to characterize highly heterogeneous ecosystems in the forest-shrubland ecotone. We analyzed a 22-year Landsat time series (1985&ndash;2012) to determine changes in forest in an area that experienced a relatively dry decade punctuated by two years of extreme drought. We assessed the relationship between several vegetation indices and field measured characteristics (e.g. plant area index and canopy gap fraction) and applied these indices to trend analysis to uncover the location, direction and timing of change. Finally, we assessed the interaction of climate and topography by forest functional type. The Normalized Difference Moisture Index (NDMI), a measure of canopy water content, had the strongest correlation with short-term field measures of plant area index ( R 2 = 0.64) and canopy gap fraction ( R 2 = 0.65). Over the entire time period, 25% of the forested area experienced a significant ( p -value < 0.05) negative trend in NDMI, compared to less than 10% in a positive trend. Coniferous forests were more likely to be associated with a negative NDMI trend than deciduous forest. Forests on southern aspects were least likely to exhibit a negative trend while north aspects were most prevalent. Field plots with a negative trend had a lower live density, and higher amounts of standing dead and down trees compared to plots with no trend. Our analysis identifies spatially explicit patterns of long-term trends anchored with ground based evidence to highlight areas of forest that are resistant, persistent or vulnerable to severe drought. The results provide a long-term perspective for the resource management of this area and can be applied to similar ecosystems throughout western North America.

Colorado, Utah, Wyoming

Carbon, oxygen and hydrogen isotope abundances inStylites reflect its unique physiology

Natural abundances of oxygen-18, carbon-14, carbon-13 and deuterium were measured in Stylites andicola , a terrestrial vascular plant lacking stomates, and two terrestrial plant species having stomates that grew nearby. No substantial differences in oxygen-18 and carbon-13 abundances were observed among the three species. The deuterium concentration in Stylites was much higher than in the other species, confirming a previous report of CAM in Stylites. Stylites was depleted in carbon-14 relative to the present day atmosphere, consistent with the proposal that it fixes CO 2 derived from decomposing peat.

Oecologia

Shifting species interactions in terrestrial dryland ecosystems under altered water availability and climate change

Species interactions play key roles in linking the responses of populations, communities, and ecosystems to environmental change. For instance, species interactions are an important determinant of the complexity of changes in trophic biomass with variation in resources. Water resources are a major driver of terrestrial ecology and climate change is expected to greatly alter the distribution of this critical resource. While previous studies have documented strong effects of global environmental change on species interactions in general, responses can vary from region to region. Dryland ecosystems occupy more than one-third of the Earth's land mass, are greatly affected by changes in water availability, and are predicted to be hotspots of climate change. Thus, it is imperative to understand the effects of environmental change on these globally significant ecosystems. Here, we review studies of the responses of population-level plant-plant, plant-herbivore, and predator-prey interactions to changes in water availability in dryland environments in order to develop new hypotheses and predictions to guide future research. To help explain patterns of interaction outcomes, we developed a conceptual model that views interaction outcomes as shifting between (1) competition and facilitation (plant-plant), (2) herbivory, neutralism, or mutualism (plant-herbivore), or (3) neutralism and predation (predator-prey), as water availability crosses physiological, behavioural, or population-density thresholds. We link our conceptual model to hypothetical scenarios of current and future water availability to make testable predictions about the influence of changes in water availability on species interactions. We also examine potential implications of our conceptual model for the relative importance of top-down effects and the linearity of patterns of change in trophic biomass with changes in water availability. Finally, we highlight key research needs and some possible broader impacts of our findings. Overall, we hope to stimulate and guide future research that links changes in water availability to patterns of species interactions and the dynamics of populations and communities in dryland ecosystems.

Biological Reviews

Surfactants as blackbird stressing agents

Applications of wetting-agent solutions produce mortality in birds. The exact cause of death is undetermined but it is believed that destruction of the insulating qualities of the plumage permits ambient cold temperatures and evaporation to lower the body temperature to a lethal level. The original concept of using these materials as bird-control tools was developed in 1958 at the Patuxent Wildlife Research Center, Bureau of Sport Fisheries and Wildlife Laurel, Maryland. Early field trials by personnel of the Division of Wildlife Services and the Denver Wildlife Research Center indicated that ground-application techniques had promise but limitations of the equipment precluded successful large-scale roost treatments. In 1966, Patuxent Center personnel began using tanker-type aircraft to evaluate high-volume aerial applications of wetting agents. The success of these tests led to the use of small aircraft to make low-volume, high-concentration aerial applications just prior to expected rainfall. Recent trials of the low-volume method show that, with some limitations, it is effective, inexpensive, and safe to the environment. Current research emphasizes the screening of new candidate materials for efficacy, biodegradability, and toxicity to plants and non-target animals, as well as basic investigations of the avian physiological mechanisms involved. Field trials to develop more effective application techniques will continue.

Proceedings of the Vertebrate Pest Conference

Is there evidence of adaptation to tidal flooding in saplings of baldcypress subjected to different salinity regimes?

Plant populations may adapt to environmental conditions over time by developing genetically based morphological or physiological characteristics. For tidal freshwater forested wetlands, we hypothesized that the conditions under which trees developed led to ecotypic difference in response of progeny to hydroperiod. Specifically, we looked for evidence of ecotypic adaptation for tidal flooding at different salinity regimes using growth and ecophysiological characteristics of two tidal and two non-tidal source collections of baldcypress (Taxodium distichum (L.) L.C. Rich) from the southeastern United States. Saplings were subjected to treatments of hydrology (permanent versus tidal flooding) and salinity (0 versus ???2 g l -1 ) for two and a half growing seasons in a greenhouse environment. Saplings from tidal sources maintained 21-41% lower overall growth and biomass accumulation than saplings from non-tidal sources, while saplings from non-tidal sources maintained 14-19% lower overall rates of net photosynthetic assimilation, leaf transpiration, and stomatal conductance than saplings from tidal sources. However, we found no evidence for growth or physiological enhancement of saplings from tidal sources to tide, or of saplings from non-tidal sources to no tide. All saplings growing under permanent flooding exhibited reduced growth and leaf gas exchange regardless of source, with little evidence for consistent salinity effects across hydroperiods. While we reject our original hypothesis, we suggest that adaptations of coastal baldcypress to broad (rather than narrow) environmental conditions may promote ecophysiological and growth enhancements under a range of global-change-induced stressors, perhaps reflecting a natural resilience to environmental change while precluding adaptations for specific flood regimes.

Environmental and Experimental Botany

Mechanisms of plant survival and mortality during drought: Why do some plants survive while others succumb to drought?

Severe droughts have been associated with regional-scale forest mortality worldwide. Climate change is expected to exacerbate regional mortality events; however, prediction remains difficult because the physiological mechanisms underlying drought survival and mortality are poorly understood. We developed a hydraulically based theory considering carbon balance and insect resistance that allowed development and examination of hypotheses regarding survival and mortality. Multiple mechanisms may cause mortality during drought. A common mechanism for plants with isohydric regulation of water status results from avoidance of drought-induced hydraulic failure via stomatal closure, resulting in carbon starvation and a cascade of downstream effects such as reduced resistance to biotic agents. Mortality by hydraulic failure per se may occur for isohydric seedlings or trees near their maximum height. Although anisohydric plants are relatively drought-tolerant, they are predisposed to hydraulic failure because they operate with narrower hydraulic safety margins during drought. Elevated temperatures should exacerbate carbon starvation and hydraulic failure. Biotic agents may amplify and be amplified by drought-induced plant stress. Wet multidecadal climate oscillations may increase plant susceptibility to drought-induced mortality by stimulating shifts in hydraulic architecture, effectively predisposing plants to water stress. Climate warming and increased frequency of extreme events will probably cause increased regional mortality episodes. Isohydric and anisohydric water potential regulation may partition species between survival and mortality, and, as such, incorporating this hydraulic framework may be effective for modeling plant survival and mortality under future climate conditions.

New Phytologist

Analysis of utilization of desert habitats with dynamic simulation

The effects of climate and herbivores on cool desert shrubs in north-western Utah were investigated with a dynamic simulation model. Cool desert shrublands are extensively managed as grazing lands, and are defoliated annually by domestic livestock. A primary production model was used to simulate harvest yields and shrub responses under a variety of climatic regimes and defoliation patterns. The model consists of six plant components, and it is based on equations of growth analysis. Plant responses were simulated under various combinations of 20 annual weather patterns and 14 defoliation strategies. Results of the simulations exhibit some unexpected linearities in model behavior, and emphasize the importance of both the pattern of climate and the level of plant vigor in determining optimal harvest strategies. Model behaviors are interpreted in terms of shrub morphology, physiology and ecology.

Journal of Environmental Management

Niche evolution and adaptive radiation: Testing the order of trait divergence

In the course of an adaptive radiation, the evolution of niche parameters is of particular interest for understanding modes of speciation and the consequences for coexistence of related species within communities. We pose a general question: In the course of an evolutionary radiation, do traits related to within-community niche differences (?? niche) evolve before or after differentiation of macrohabitat affinity or climatic tolerances (?? niche)? Here we introduce a new test to address this question, based on a modification of the method of independent contrasts. The divergence order test (DOT) is based on the average age of the nodes on a tree, weighted by the absolute magnitude of the contrast at each node for a particular trait. The comparison of these weighted averages reveals whether large divergences for one trait have occurred earlier or later in the course of diversification, relative to a second trait; significance is determined by bootstrapping from maximum-likelihood ancestral state reconstructions. The method is applied to the evolution of Ceanothus, a woody plant group in California, in which co-occurring species exhibit significant differences in a key leaf trait (specific leaf area) associated with contrasting physiological and life history strategies. Co-occurring species differ more for this trait than expected under a null model of community assembly. This ?? niche difference evolved early in the divergence of two major subclades within Ceanothus, whereas climatic distributions (?? niche traits) diversified later within each of the subclades. However, rapid evolution of climate parameters makes inferences of early divergence events highly uncertain, and differentiation of the ?? niche might have taken place throughout the evolution of the group, without leaving a clear phylogenetic signal. Similar patterns observed in several plant and animal groups suggest that early divergence of ?? niche traits might be a common feature of niche evolution in many adaptive radiations. ?? 2006 by the Ecological Society of America.

Ecology

Early growth and ecophysiological responses of Koa (Acacia koa A. Gray) seedlings to reduced water and phosphorus

Sites in need of restoration typically have one or more environmental factors that limit seedling establishment. Identifying ecophysiological responses to environmental stressors can provide important insights into mitigating measures that would allow seedlings to overcome such constraints to survival. Koa ( Acacia koa A. Gray) is a nitrogen-fixing tree species endemic to Hawaiʻi that is highly valued in restoring degraded forest ecosystems, which are often limited in available water and phosphorus. This study examined how koa seedlings respond to conditions of reduced water (65 W) and no phosphorus (0P). After 17 weeks, seedlings subjected to 65 W or 0P accumulated less biomass, smaller root-collar diameters, and lower nitrogen and phosphorus contents. Combined reductions in water and P resulted in seedlings with increased root to shoot dry biomass and shorter shoots. Seedlings subjected to 65 W also had lower instantaneous rates of CO 2 assimilation, but higher instantaneous water-use efficiencies following irrigation, suggesting that koa responds to water deficits by decreasing water loss via reduced stomatal conductance. Seedlings subjected to 0P had similar rates of CO 2 assimilation relative to those grown with adequate P, suggesting that koa is able to employ strategies to avoid physiological impairment from conditions of inadequate P. Future research should assess whether subjecting koa seedlings to reduced water before planting on water-limited sites cues increased drought resistance and whether uptake and storage of P by seedlings in the nursery better supports growth following outplanting, particularly on sites with anticipated low plant-available water.

Hawai'i

Soil microbial community profiles and functional diversity in limestone cedar glades

Rock outcrop ecosystems, such as limestone cedar glades (LCGs), are known for their rare and endemic plant species adapted to high levels of abiotic stress. Soils in LCGs are thin (< 25 cm), soil-moisture conditions fluctuate seasonally between xeric and saturated, and summer soil temperatures commonly exceed 48 &deg;C. The effects of these stressors on soil microbial communities (SMC) remain largely unstudied, despite the importance of SMC-plant interactions in regulating the structure and function of terrestrial ecosystems. SMC profiles and functional diversity were characterized in LCGs using community level physiological profiling (CLPP) and plate-dilution frequency assays (PDFA). Most-probable number (MPN) estimates and microbial substrate-utilization diversity (H) were positively related to soil thickness, soil organic matter (OM), soil water content, and vegetation density, and were diminished in alkaline soil relative to circumneutral soil. Soil nitrate showed no relationship to SMCs, suggesting lack of N-limitation. Canonical correlation analysis indicated strong correlations between microbial CLPP patterns and several physical and chemical properties of soil, primarily temperature at the ground surface and at 4-cm depth, and secondarily soil-water content, enabling differentiation by season. Thus, it was demonstrated that several well-described abiotic determinants of plant community structure in this ecosystem are also reflected in SMC profiles.

Tennessee

Editorial: Plant phenology shifts and their ecological and climatic consequences

Climate change is causing plant phenology to shift, with consequences for ecosystems and the Earth’s climate. Over the last decades, the timing of many important phenological events has advanced in spring, such as leaf emergence and flowering, or been delayed in fall, such as leaf coloration and leaf fall. The consequences of such phenological shifts are still largely unknown, but are hypothesized to have cascading effects on ecosystems, carbon and water cycles, and Earths’ climate. With this research topic, we aimed to synthesize and inspire innovative research in plant phenology to address research questions and challenges on the consequences of phenological shifts on ecosystem function and local hydrology. The articles presented here improve our understanding of the physiological mechanisms responsible for the current phenological changes in spring and fall and provide insight into some of the consequences of these changes on hydrological cycles and ecosystem functioning.

Frontiers in Plant Science

Population differentiation along a flood frequency gradient: Physiological adaptations to flooding in Nyssa sylvatica

Throughout the southeastern United States the hardwood Nyssa sylvatica (sensu lato) is distributed along a soil moisture gradient from upland sites, which are never flooded, to floodplains, which are periodically flooded and drained to permanently flooded swamps. Population differentiation with respect to flood tolerance and related physiological attributes was investigated using 1—year—old seedlings grown in a greenhouse from seed collected along this gradient. Upland plants were very intolerant of flooded soils. Their root systems deteriorated, root respiration rates dropped and, after a year under such conditions, survival was poor and those that did remain were greatly stunted and had accumulated large concentrations of many nutrient elements. In contrast swamp plants were quite tolerant of flooded soils. Upon flooding, parts of the orginal root system were lost but new roots were initiated which had an increased capacity for alcoholic fermentation. Many of these new roots were more succulent, larger in diameter, and less branched than drained roots. Such succulent roots however were only a temporary response to short—term flooding; plants flooded for a year did not have such roots, rather the root system superficially resembled drained roots. Concomitant with this return to drained—like roots was an increase in internal oxygen transport to the roots and a drop in alcoholic fermentation. Floodplain plants under drained conditions allocated less biomass to roots than to shoots and had high respiration rates, traits similar to upland plants. Under flooded conditions they initiated new roots with medium respiration rates, allocated less biomass to roots than to shoots, significantly increased oxygen transport to the roots and had high survival, traits similar to swamp plants. Thus, the floodplain population produced a distinctly flood—tolerant phenotype; but not nearly as tolerant of flooded conditions as the swamp phenotype. Floodplain plants differed from swamp plants in transporting less oxygen to the roots under drained conditions, initiating fewer succulent—type roots and not accelerating alcoholic fermentation upon flooding and after a year under flooded conditions having less total biomass, less oxygen transport to the roots and a greater accumulation of Fe and Mn in the roots. The floodplain plants apparently have been selected to be similar to upland plants under drained conditions and swamp plants under flooding and one consequence of this is that their tolerance of flooded conditions is intermediate. It is argued that one of the more important trade—offs in adapting to flooded conditions is that high internal oxygen transport carries with it a ‘cost’ in terms of excessive water loss under water stress conditions.

Ecological Monographs

Sensitivity of desert cryptograms to air pollutants: soil crusts and rock lichens

Parks throughout the West are being faced with increasing air pollution threats from current or proposed industries near their boundaries. For this reason, it is important to understand the effects these industries may have on desert ecosystems. Rock lichens can be excellent biomonitors, acting as early warning systems of impending damage to other components of the desert ecosystem. Cryptogamic crusts, consisting mostly of cyanobacteria and lichens, may not only be excellent bioindicators, but also are an essential part of the desert ecosystem. Their presence is critical for soil stability as well as for the contribution of nitrogen to the ecosystem in a form available to higher plants. Air pollutants, such as emissions from coal-fired power plants, may threaten the healthy functioning of these non-vascular plants. The purpose of this study is to determine if, in fact, air pollutants do have an impact on the physiological functioning of cryptogamic crusts or rock lichens in desert systems and, if so, to what extent. Some results have already been obtained. Both rock lichens and cryptogamic crusts exhibit physiological damage in the vicinity of the Navajo Generating Station in Page, Arizona. Increased electrolyte leakage and chlorophyll degradation, along with reduced nitrogen fixation, have been found. Preliminary studies comparing sensitivity between substrates indicate that crusts on limestone and sandstone substrates may be more sensitive than those on gypsum.

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