Could vesicular transport of Na+ and Cl- be a feature of salt tolerance in halophytes?

Background: Halophytes tolerate external salt concentrations of 200 mm and more, accumulating salt concentrations of 500 mm and more in their shoots; some, recretohalophytes, excrete salt through glands on their leaves. Ions are accumulated in central vacuoles, but the pathway taken by these ions from the outside of the roots to the vacuoles inside the cells is poorly understood. Do the ions cross membranes through ion channels and transporters or move in vesicles, or both? Vesicular transport from the plasma membrane to the vacuole would explain how halophytes avoid the toxicity of high salt concentrations on metabolism. There is also a role for vesicles in the export of ions via salt glands. Scope and Methods: We have collected data on the fluxes of sodium and chloride ions in halophytes, based on the weight of the transporting organs and on the membrane area across which the flux occurs; the latter range from 17 nmol m-2 s-1 to 4.2 µmol m-2 s-1 and values up to 1 µmol m-2 s-1 need to be consistent with whatever transport system is in operation. We have summarized the sizes and rates of turnover of vesicles in plants, where clathrin-independent vesicles are 100 nm or more in diameter and can merge with the plasma membrane at rates of 100 s-1. We gathered evidence for vesicular transport of ions in halophytes and evaluated whether vesicular transport could account for the observable fluxes. Conclusions: There is strong evidence in favour of vesicular transport in plants and circumstantial evidence in favour of the movement of ions in vesicles. Estimated rates of vesicle turnover could account for ion transport at the lower reported fluxes (around 20 nmol m-2 s-1), but the higher fluxes may require vesicles of the order of 1 µm or more in diameter. The very high fluxes reported in some salt glands might be an artefact of the way they were measured.

Application and Comparison of the MODIS-Derived Enhanced Vegetation Index to VIIRS, Landsat 5 TM and Landsat 8 OLI Platforms: A Case Study in the Arid Colorado River Delta, Mexico.

The Enhanced Vegetation Index (EVI) is a key Earth science parameter used to assess vegetation, originally developed and calibrated for the Moderate Resolution Imaging Spectroradiometer (MODIS) aboard the Terra and Aqua satellites. With the impending decommissioning of the MODIS sensors by the year 2020/2022, alternative platforms will need to be used to estimate EVI. We compared Landsat 5 (2000⁻2011), 8 (2013⁻2016) and the Visible Infrared Imaging Radiometer Suite (VIIRS; 2013⁻2016) to MODIS EVI (2000⁻2016) over a 420,083-ha area of the arid lower Colorado River Delta in Mexico. Over large areas with mixed land cover or agricultural fields, we found high correspondence between Landsat and MODIS EVI (R² = 0.93 for the entire area studied and 0.97 for agricultural fields), but the relationship was weak over bare soil (R² = 0.27) and riparian vegetation (R² = 0.48). The correlation between MODIS and Landsat EVI was higher over large, homogeneous areas and was generally lower in narrow riparian areas. VIIRS and MODIS EVI were highly similar (R² = 0.99 for the entire area studied) and did not show the same decrease in performance in smaller, narrower regions as Landsat. Landsat and VIIRS provide EVI estimates of similar quality and characteristics to MODIS, but scale, seasonality and land cover type(s) should be considered before implementing Landsat EVI in a particular area.

Can local adaptation explain varying patterns of herbivory tolerance in a recently introduced woody plant in North America?

Patterns of woody-plant mortality have been linked to global-scale environmental changes, such as extreme drought, heat stress, more frequent and intense fires, and episodic outbreaks of insects and pathogens. Although many studies have focussed on survival and mortality in response to specific physiological stresses, little attention has been paid to the role of genetic heritability of traits and local adaptation in influencing patterns of plant mortality, especially in non-native species. Tamarix spp. is a dominant, non-native riparian tree in western North America that is experiencing dieback in some areas of its range due to episodic herbivory by the recently introduced northern tamarisk leaf beetle (Diorhabda carinulata). We propose that genotype × environment interactions largely underpin current and future patterns of Tamarix mortality. We anticipate that (i) despite its recent introduction, and the potential for significant gene flow, Tamarix in western North America is generally adapted to local environmental conditions across its current range in part due to hybridization of two species; (ii) local adaptation to specific climate, soil and resource availability will yield predictable responses to episodic herbivory; and (iii) the ability to cope with a combination of episodic herbivory and increased aridity associated with climate change will be largely based on functional tradeoffs in resource allocation. This review focusses on the potential heritability of plant carbon allocation patterns in Tamarix, focussing on the relative contribution of acquired carbon to non-structural carbohydrate (NSC) pools versus other sinks as the basis for surviving episodic disturbance. Where high aridity and/or poor edaphic position lead to chronic stress, NSC pools may fall below a minimum threshold because of an imbalance between the supply of carbon and its demand by various sinks. Identifying patterns of local adaptation of traits related to resource allocation will improve forecasting of Tamarix population susceptibility to episodic herbivory.

Effects of grazing on leaf area index, fractional cover and evapotranspiration by a desert phreatophyte community at a former uranium mill site on the Colorado Plateau.

This study employed ground and remote sensing methods to monitor the effects of grazing on leaf area index (LAI), fractional cover (f(c)) and evapotranspiration (ET) of a desert phreatophyte community over an 11 year period at a former uranium mill site on the Colorado Plateau, U.S. Nitrate, ammonium and sulfate are migrating away from the mill site in a shallow alluvial aquifer. The phreatophyte community, consisting of Atriplex canescens (ATCA) and Sarcobatus vermiculatus (SAVE) shrubs, intercepts groundwater and could potentially slow the movement of the contaminant plume through evapotranspiration (ET). However, the site has been heavily grazed by livestock, reducing plant cover and LAI. We used livestock exclosures and revegetation plots to determine the effects of grazing on LAI, f(c) and ET, then projected the findings over the whole site using multi-platform remote sensing methods. We show that ET is approximately equal to annual precipitation at the site, but when ATCA and SAVE are protected from grazing they can develop high f(c) and LAI values, and ET can exceed annual precipitation, with the excess coming from groundwater discharge. Therefore, control of grazing could be an effective method to slow migration of contaminants at this and similar sites in the western U.S.

The effect of wetland vegetation on the survival of Escherichia coli, Salmonella typhimurium, bacteriophage MS-2 and polio virus.

A study was conducted to examine the role of aquatic plants used in constructed wetlands on the survival of enteric bacteria and viruses. Four small-scale wetland systems, receiving fresh water and two other wetland systems, receiving secondary unchlorinated sewage were used in this study. Fresh water and secondary sewage without the presence of any aquatic plants were used as controls. Escherichia coli, Salmonella typhimurium, bacteriophage MS-2 and poliovirus were added to the waters collected from the wetlands and controls. The presence of aquatic plants significantly increased the die-off of both bacteria in fresh water and secondary sewage. No significant difference in the die-off of E. coli and S. typhimurium was observed in water from wetlands with different types of plants in freshwater. However, there was a significant difference in the die-off of E. coli in water with aquatic plants when sewage was used. The presence of the plants significantly increased the inactivation of MS-2 and poliovirus. Additional work on the survival of E. coli indicated that the plausible mechanism of bacterial die-off in constructed wetlands is through increased microbial competition or predation.

Reconciling environmental and flood control goals on an arid-zone river: case study of the limitrophe region of the lower colorado river in the United States and Mexico.

Arid zone rivers have highly variable flow rates, and flood control projects are needed to protect adjacent property from flood damage. On the other hand, riparian corridors provide important wildlife habitat, especially for birds, and riparian vegetation is adapted to the natural variability in flows on these rivers. While environmental and flood control goals might appear to be at odds, we show that both goals can be accommodated in the Limitrophe Region (the shared border between the United States and Mexico) on the Lower Colorado River. In 1999, the International Boundary and Water Commission proposed a routine maintenance project to clear vegetation and create a pilot channel within the Limitrophe Region to improve flow capacity and delineate the border. In 2000, however, Minute 306 to the international water treaty was adopted, which calls for consideration of environmental effects of IBWC actions. We conducted vegetation and bird surveys within the Limitrophe and found that this river segment is unusually rich in native cottonwood and willow trees, marsh habitat, and resident and migratory birds compared to flow-regulated segments of river. A flood-frequency analysis showed that the existing levee system can easily contain a 100 year flood even if vegetation is not removed, and the existing braided channel system has greater carrying capacity than the proposed pilot channel.

Relationship Between Remotely-sensed Vegetation Indices, Canopy Attributes and Plant Physiological Processes: What Vegetation Indices Can and Cannot Tell Us About the Landscape.

Vegetation indices (VIs) are among the oldest tools in remote sensing studies. Although many variations exist, most of them ratio the reflection of light in the red and NIR sections of the spectrum to separate the landscape into water, soil, and vegetation. Theoretical analyses and field studies have shown that VIs are near-linearly related to photosynthetically active radiation absorbed by a plant canopy, and therefore to light-dependent physiological processes, such as photosynthesis, occurring in the upper canopy. Practical studies have used time-series VIs to measure primary production and evapotranspiration, but these are limited in accuracy to that of the data used in ground truthing or calibrating the models used. VIs are also used to estimate a wide variety of other canopy attributes that are used in Soil-Vegetation-Atmosphere Transfer (SVAT), Surface Energy Balance (SEB), and Global Climate Models (GCM). These attributes include fractional vegetation cover, leaf area index, roughness lengths for turbulent transfer, emissivity and albedo. However, VIs often exhibit only moderate, non-linear relationships to these canopy attributes, compromising the accuracy of the models. We use case studies to illustrate the use and misuse of VIs, and argue for using VIs most simply as a measurement of canopy light absorption rather than as a surrogate for detailed features of canopy architecture. Used this way, VIs are compatible with "Big Leaf" SVAT and GCMs that assume that canopy carbon and moisture fluxes have the same relative response to the environment as any single leaf, simplifying the task of modeling complex landscapes.

Riparian vegetation dynamics and evapotranspiration in the riparian corridor in the delta of the Colorado River, Mexico.

Like other great desert rivers, the Colorado River in the United States and Mexico is highly regulated to provide water for human use. No water is officially allotted to support the natural ecosystems in the delta of the river in Mexico. However, precipitation is inherently variable in this watershed, and from 1981-2004, 15% of the mean annual flow of the Lower Colorado River has entered the riparian corridor below the last diversion point for water in Mexico. These flows include flood releases from US dams and much smaller administrative spills released back to the river from irrigators in the US and Mexico. These flows have germinated new cohorts of native cottonwood and willow trees and have established an active aquatic ecosystem in the riparian corridor in Mexico. We used ground and remote-sensing methods to determine the composition and fractional cover of the vegetation in the riparian corridor, its annual water consumption, and the sources of water that support the ecosystem. The study covered the period 2000-2004, a flood year followed by 4 dry years. The riparian corridor occupies 30,000 ha between flood control levees in Mexico. Annual evapotranspiration (ET), estimated by Moderate Resolution Imaging Spectrometer (MODIS) satellite imagery calibrated against moisture flux tower data, was about 1.1 m yr(-1) and was fairly constant throughout the study period despite a paucity of surface flows 2001-2004. Total ET averaged 3.4 x 10(8)m(3)yr(-1), about 15% of Colorado River water entering Mexico from the US Surface flows could have played only a small part in supporting these high ET losses. We conclude that the riparian ET is supported mainly by the shallow regional aquifer, derived from agricultural return flows, that approaches the surface in the riparian zone. Nevertheless, surface flows are important in germinating cohorts of native trees, in washing salts from the soil and aquifer, and in providing aquatic habitat, thereby enriching the habitat value of the riparian corridor for birds and other wildlife. Conservation and water management strategies to enhance the delta habitats are discussed in light of the findings.

Bacterial community changes during plant establishment at the San Pedro River mine tailings site.

Mine tailings are moderately to severely impacted sites that lack normal plant cover, soil structure and development, and the associated microbial community. In arid and semiarid environments, tailings and their associated contaminants are prone to eolian dispersion and water erosion, thus becoming sources of metal contamination. One approach to minimize or eliminate these processes is to establish a permanent vegetation cover on tailings piles. Here we report a revegetation trial conducted at a moderately impacted mine tailings site in southern Arizona. A salt and drought-tolerant plant, four-wing saltbush [Atriplex canescens (Pursh) Nutt.], was chosen for the trial. A series of 3 by 3 m plots were established in quadruplicate on the test site to evaluate growth of four-wing saltbush transplants alone or with compost addition. Results show that >80% of the transplanted saltbush survived after 1.5 yr in both treatments. Enumeration of heterotrophs and community structure analysis were conducted to monitor bacterial community changes during plant establishment as an indicator of plant and soil health. The bacterial community was evaluated using denaturing gradient gel electrophoresis (DGGE) analysis of 16S rDNA PCR gene products from tailings samples taken beneath transplant canopies. Significant differences in heterotrophic counts and community composition were observed between the two treatments and unplanted controls throughout the trial, but treatment effects were not observed. The results suggest that compost is not necessary for plant establishment at this site and that plants, rather than added compost, is the primary factor enhancing bacterial heterotrophic counts and affecting community composition.

Just add water and the Colorado River still reaches the sea.

A recent article in Environmental Management by All argued that flood flows in North America's Colorado River do not reach the Gulf of California because they are captured and evaporated in Laguna Salada, a below sea-level lakebed near the mouth of the river. We refute this hypothesis by showing that (1) due to its limited area, the Laguna Salada could have evaporated less than 10% of the flood flows that have occurred since 1989; (2) low flow volumes preferentially flow to the Gulf rather than Laguna Salada; (3) All's method for detecting water surface area in the Laguna Salada appears to be flawed because Landsat Thematic Mapper images of the lakebed show it to be dry when All's analyses said it was flooded; (4) direct measurements of salinity at the mouth of the river and in the Upper Gulf of California during flood flows in 1993 and 1998 confirm that flood waters reach the sea; and (5) stable oxygen isotope signatures in clam shells and fish otoliths recorded the dilution of seawater with fresh water during the 1993 and 1998 flows. Furthermore, All's conclusion that freshwater flows do not benefit the ecology of the marine zone is incorrect because the peer-reviewed literature shows that postlarval larval shrimp populations increase during floods, and the subsequent year's shrimp harvest increases. Furthermore, freshwater flows increase the nursery area for Gulf corvina (Cynoscion othonopterus), an important commercial fish that requires estuarine habitats with salinities in the range of 26-38 per thousand during its natal stages. Although flood flows are now much diminished compared to the pre-dam era, they are still important to the remnant wetland and riparian habitats of the Colorado River delta and to organisms in the intertidal and marine zone. Only a small fraction of the flood flows are evaporated in Laguna Salada.

Phytostabilization potential of quailbush for mine tailings: growth, metal accumulation, and microbial community changes.

Abandoned mine tailings sites in semiarid regions remain unvegetated for extended periods of time and are subject to eolian dispersion and water erosion. This study examines the potential phytostabilization of a lead-zinc mine tailings site using a native, drought-tolerant halophyte, quailbush [Atriplex lentiformis (Torr.) S. Wats.]. In a greenhouse study germination, growth, and metal uptake was evaluated in two compost-amended mine tailings samples, K4 (pH 3) and K6 (pH 6) at 75, 85, 90, 95, and 100% mine tailings, and two controls, off-site and compost. Microbial community changes were monitored by performing MPN analysis of iron- and sulfur-oxidizing bacteria as well as heterotrophic plate counts. Results demonstrate that germination is not a good indicator for phytostabilization since it was only inhibited in the unamended K4 treatment. Plant growth was significantly reduced in 95 and 100% mine tailings, while growth in 75, 85, and 90% treatments was similar to the off-site control. Quailbush accumulated elevated levels of the nutrient metals Na, K, Mn, and Zn in the shoot tissues; however, metal accumulation was generally below the domestic animal toxicity limit. Initially, autotrophic population estimates were four to six logs higher than heterotrophic counts, indicating extremely stressed conditions. However, post-harvest, heterotrophic bacterial counts increased to normal levels (approximately 10(6) CFU g-1 dry tailings) and dominated the rhizosphere. Therefore, with compost amendment, quailbush has good potential as a native species candidate for phytostabilization of mine tailings in semiarid environments.

Salt tolerance and osmotic adjustment of Spartina alterniflora (Poaceae) and the invasive M haplotype of Phragmites australis (Poaceae) along a salinity gradient.

An invasive variety of Phragmites australis (Poaceae, common reed), the M haplotype, has been implicated in the spread of this species into North American salt marshes that are normally dominated by the salt marsh grass Spartina alterniflora (Poaceae, smooth cordgrass). In some European marshes, on the other hand, Spartina spp. derived from S. alterniflora have spread into brackish P. australis marshes. In both cases, the non-native grass is thought to degrade the habitat value of the marsh for wildlife, and it is important to understand the physiological processes that lead to these species replacements. We compared the growth, salt tolerance, and osmotic adjustment of M haplotype P. australis and S. alterniflora along a salinity gradient in greenhouse experiments. Spartina alterniflora produced new biomass up to 0.6 M NaCl, whereas P. australis did not grow well above 0.2 M NaCl. The greater salt tolerance of S. alterniflora compared with P. australis was due to its ability to use Na(+) for osmotic adjustment in the shoots. On the other hand, at low salinities P. australis produced more shoots per gram of rhizome tissue than did S. alterniflora. This study illustrates how ecophysiological differences can shift the competitive advantage from one species to another along a stress gradient. Phragmites australis is spreading into North American coastal marshes that are experiencing reduced salinities, while Spartina spp. are spreading into northern European brackish marshes that are experiencing increased salinities as land use patterns change on the two continents.

Vegetation mapping for change detection on an arid-zone river.

A vegetation mapping system for change detection was tested at the Havasu National Wildlife Refuge (HNWR) on the Lower Colorado River. A low-cost, aerial photomosaic of the 4200 ha, study area was constructed utilizing an automated digital camera system, supplemented with oblique photographs to aid in determining species composition and plant heights. Ground-truth plots showed high accuracy in distinguishing native cottonwood (Populus fremontii) and willow (Salix gooddingii) trees from other vegetation on aerial photos. Marsh vegetation (mainly cattails, Typha domengensis) was also easily identified. However, shrubby terrestrial vegetation, consisting of saltcedar (Tamarix ramosissima), arrowweed (Pluchea sericea), and mesquite trees (Prosopis spp.), could not be accurately distinguished from each other and were combined into a single shrub layer on the final vegetation map. The final map took the form of a base, shrub and marsh layer, which was displayed as a Normalized Difference Vegetation Index map from a Landsat Enhanced Thematic Mapper (ETM+) image to show vegetation intensity. Native willow and cottonwood trees were digitized manually on the photomosaic and overlain on the shrub layer in a GIS. By contrast to present, qualitative mapping systems used on the Lower Colorado River, this mapping system provides quantitative information that can be used for accurate change detection. However, better methods to distinguish between saltcedar, mesquite, and arrowweed are needed to map the shrub layer.

A comparison of chelator-facilitated metal uptake by a halophyte and a glycophyte.

Phytoextraction is the use of plants to remove contaminants, in particular metals, from soil via root uptake and translocation to the shoots. Efficient phytoextraction requires high-biomass plants with efficient translocating properties. Halophytes characteristically accumulate large quantities of salts in above ground tissue material and can have high biomass production. It has been speculated that salt-tolerant plants may also be heavy metal tolerant and, further, may be able to accumulate metals. This study compared growth and metal uptake by a halophyte, Atriplex nummularia, and a common glycophyte, Zea mays, in a mine-tailing contaminated soil:mulch mixture. Two chelators, ethylenediaminetetraacetic acid (EDTA) and rhamnolipid, were used to facilitate plant metal uptake. Despite a lower growth rate (2% growth/d) in the contaminated soil, the halophyte accumulated roughly the same amount of metals as the glycophyte on a mass basis (30-40 mg/kg dry wt). Neither plant, however, hyperaccumulated any of the metals tested. When treated with EDTA, specific differences in patterns of metal uptake between the two plants emerged. The halophyte accumulated significantly more Cu (2x) and Pb (1x) in the shoots than the glycophyte, but root metal concentrations were generally higher for the glycophyte, indicating that the halophyte translocated more metal from the root to the shoot than the glycophyte. For example, Zn shoot-to-root ratios ranged from 1.4 to 2.1 for Atriplex and from 0.5 to 0.6 for Z. mays. The biodegradable chelator rhamnolipid was not effective at enhancing shoot metal concentrations, even though radiolabeled chelator was found in the shoot material of both plants. Our results suggest that halophytes, despite their slower growth rates, may have greater potential to selectively phytoextract metals from contaminated soils than glycophytes.

Microbiology of Hydroponically-Grown Lettuce.

The microbial quality of lettuce ( Lactuca sativa L. var. "Ostinata") cultivated in a hydroponic system was evaluated. Over a 3-month study period, samples of lettuce, nutrient solution, and peat-vermiculite growing mixture from the greenhouse were analyzed for total aerobic bacteria, yeasts, molds, and coliforms. There was a consistent amount of each type of organism occurring within each sample group for a given month, and the numbers of aerobic bacteria and coliforms present were generally similar to those reported for lettuce and leafy vegetable crops propagated by the conventional method of agriculture. Over the study period, the modal values for each type of organism in lettuce were: aerobic bacteria, 7.9 × 106 CFU/g (range of 3.8 × 104 to 2.3 × 108); coliforms, 1.5 × 104 CFU/g (range of none detected to greater than or equal to 5.3 × 107); molds, 2.9 × 103 CFU/g (range of 1.2 × 102 to 5.3 × 104); and yeasts, 2.4 × 104 CFU/g (range of 6.9 × 102 to 2.3 × 106). The primary organisms associated with the growing system were Citrobacter freundii , Enterobacter cloacae , and Enterobacter agglomerans . No organisms of human health concern (i.e. Salmonella spp., Clostrium botulinum , Escherichia coli , or Staphylococcus aureus ) were detected in the samples. The bacteriology of lettuce produced for market by this type of hydroponic farming and packaging appears to be generally comparable to that of field-grown lettuce and to present no unique microbiological hazards to consumers.