Development of an epiphyte indicator of nutrient enrichment: A critical evaluation of observational and experimental studies

An extensive review of the literature describing epiphytes on submerged aquatic vegetation (SAV), especially seagrasses, was conducted in order to evaluate the evidence for response of epiphyte metrics to increased nutrients. Evidence from field observational studies, together with laboratory and field mesocosm experiments, was assembled from the literature and evaluated for a hypothesized positive response to nutrient addition. There was general consistency in the results to confirm that elevated nutrients tended to increase the load of epiphytes on the surface of SAV, in the absence of other limiting factors. In spite of multiple sources of uncontrolled variation, positive relationships of epiphyte load to nutrient concentration or load (either nitrogen or phosphorus) often were observed along strong anthropogenic or natural nutrient gradients in coastal regions. Such response patterns may only be evident for parts of the year. Results from both mesocosm and field experiments also generally support the increase of epiphytes with increased nutrients, although outcomes from field experiments tended to be more variable. Relatively few studies with nutrient addition in mesocosms have been done with tropical or subtropical species, and more such controlled experiments would be helpful. Experimental duration influenced results, with more positive responses of epiphytes to nutrients at shorter durations in mesocosm experiments versus more positive responses at longer durations in field experiments. In the field, response of epiphyte biomass to nutrient additions was independent of climate zone. Mesograzer activity was a critical covariate for epiphyte response under experimental nutrient elevation, but the epiphyte response was highly dependent on factors such as grazer identity and density, as well as nutrient and ambient light levels. The balance of evidence suggests that epiphytes on SAV will be a useful indicator of persistent nutrient enhancement in many situations. Careful selection of appropriate temporal and spatial constraints for data collection, and concurrent evaluation of confounding factors will help increase the signal to noise ratio for this indicator.     

Interaction between atmospheric and pedospheric nitrogen nutrition in spruce (Picea abies L. Karst) seedlings

The effect of NO₂ fumigation on root N uptake and metabolism was investigated in 3-month-old spruce (Picea abics L. Karst) seedlings. In a first experiment, the contribution of NO₂ to the plant N budget was measured during a 48 h fumigation with 100mm³m^?3 NO₂. Plants were pre-treated with various nutrient solutions containing NO₂ and NH₄⁺, NO₃^? only or no nitrogen source for 1 week prior to the beginning of fumigation. Absence of NH₄⁺ in the solution for 6d led to an increased capacity for NO₃^? uptake, whereas the absence of both ions caused a decrease in the plant N concentration, with no change in NO₃^? uptake. In fumigated plants, NO₂ uptake accounted for 20–40% of NO₃^? uptake. Root NO₃^? uptake in plants supplied with NH₄⁺plus NO₃^? solutions was decreased by NO₂ fumigation, whereas it was not significantly altered in the other treatments. In a second experiment, spruce seedlings were grown on a solution containing both NO₂ and NH₄⁺ and were fumigated or not with 100mm³m^?3 NO₂ for 7 weeks. Fumigated plants accumulated less dry matter, especially in the roots. Fluxes of the two N species were estimated from their accumulations in shoots and roots, xylem exudate analysis and ¹⁵N labelling. Root NH₄⁺ uptake was approximately three times higher than NO₃^? uptake. Nitrogen dioxide uptake represented 10–15% of the total N budget of the plants. In control plants, N assimilation occurred mainly in the roots and organic nitrogen was the main form of N transported to the shoot. Phloem transport of organic nitrogen accounted for 17% of its xylem transport. In fumigated plants, neither NO₃^? nor NH₄⁺ accumulated in the shoot, showing that all the absorbed NO₂ was assimilated. Root NO₃^? reduction was reduced whereas organic nitrogen transport in the phloem increased by a factor of 3 in NO₂-fimugated as compared with control plants. The significance of the results for the regulation of whole-plant N utilization is discussed.     

Genetics of Azospirillum brasilense with respect to ammonium transport,sugar uptake,and chemotaxis

This paper describes molecular aspects of Azospirillum-plant root association with respect to nitrogen flux and carbon utilization. In the first part, biochemical and genetic data are reported on the transport of ammonium and methylammonium in A. brasilense cells. Ammonium excreting A. brasilense mutants reported so far appear to result from alterations in genes encoding for enzymes involved in ammonium assimilation. Solid genetic evidence is given on the occurrence of a postulated ammonium transporter in A. brasilense. In the second part, biochemical and genetic evidence is likewise given for the occurrence of a high-affinity uptake system for D-galactose in A. brasilense. A sugar- binding protein that is part of this uptake system is required for chemotaxis of A. brasilense towards particular sugars, including D-galactose.     

Dynamics of concentrations and nutrient fluxes in the xylem of Ricinus communis– diurnal course, impact of nutrient availability and nutrient uptake

Diurnal courses of nutrient transport in the xylem and their response to external availability of nutrients were studied. In soil culture, maximal concentrations in all analysed substances were observed during night‐time. Over experimental periods of up to 20 d, concentrations of some ions increased, most by accumulation in the soil. Stringent nutrient conditions were established in a novel pressure chamber. An aeroponic nutrient delivery system inside allows the sampling of xylem sap from intact plants under full control of the nutrient conditions at the root. Analysis of xylem transport under these highly defined conditions established that (1) diurnal variations in concentrations and fluxes in the xylem are dominated by plant‐internal processes; (2) concentrations of nutrients in the xylem sap are highly but specifically correlated with each other; (3) nitrate uptake and nitrate flux to the shoot are largely uncoupled; and (4) in continuous light, diurnal variations of xylem sap concentrations vanish. Step changes in nitrate concentrations of the nutrient solution established that (5) the concomitant increase in nitrate concentration and flux in the xylem is delayed by 2–3 h and is only transient. Diurnal variations of xylem sap composition and use of the new technique to elucidate xylem‐transport mechanisms are discussed.     

Nitrate induced iron deficiency chlorosis in Juncus acutiflorus

Chlorosis caused by iron deficiency is commonly associated with high bicarbonate levels in the soil. However, in rare cases such chlorosis has been observed in soils with high nitrate levels. In a dutch rich-fen, chlorosis has been noted in stands of Juncus acutiflorus at locations where groundwater containing high levels of nitrate reached the surface. Experiments revealed that the chlorosis could be attributed to iron deficiency although iron levels in the shoots were well above the known physiological threshold values for iron deficiency. It is postulated that increased nitrate assimilation leads to an increased apoplastic pH and to a concomitant immobilisation of iron and/or lower iron (III) reduction. Moreover free amino acid levels were markedly higher in the iron deficient plants in the field. It was found, however, that the percentage of nitrogen present as free amino acids was not influenced directly by low iron levels but mainly by the C/N ratios in the shoots. Nowadays, nitrate concentrations in ground water as high 1000 µM are no longer an exception in the Netherlands. We propose that strongly increased nitrate inputs may cause iron stress in natural vegetations, especially in wet habitats.     

Streamflow-induced variations in nitrate flux in tributaries to the Atlantic Coastal Zone

Streamflow-related variability in nutrient flux represents an important source of uncertainty in managing nutrient inputs to coastal ecosystems. Quantification of flux variability is of particular interest to coastal resource managers in adopting effective nutrient-reduction goals and monitoring progress towards these goals. We used historical records of streamflow and water-quality measurements for 104 river monitoring stations in an analysis of variability in annual and seasonal flux of nitrate to the Atlantic coastal zone. We present two measures of temporal flux variability: the coefficient of variation (CV) and the exceedence probability (EP) of 1.5 times the median flux. The magnitude of flux variations spans a very wide range and depends importantly upon the season of year and the climatic and land-use characteristics of the tributary watersheds. Year-to-year variations (CV) in annual mean flux range over two orders of magnitude, from 3–200% of the long-term mean flux, although variations more typically range from 20–40% of the long-term mean. The annual probability of exceeding the long-term median flux by more than 50% (EP) is less than 0.10 in most rivers, but is between 0.10 and 0.35 in 40% of the rivers. Year-to-year variability in seasonal mean flux commonly exceeds that in annual flux by a factor of 1.5 to 4. In western Gulf of Mexico coastal rivers, the year-to-year variablity in the seasonal mean flux is larger than in other regions, and is of a similar magnitude in all seasons. By contrast, in Atlantic coastal rivers, the winter and spring seasons, which account for about 70% of the annual flux, display the smallest relative variability in seasonal mean flux. We quantify the elasticity of nutrient flux to hypothetical changes in Streamflow (i.e., the percent increase in flux per percentage increase in mean discharge) to allow the approximation of flux variability from streamflow records and the estimation of the effects of future climatically-induced changes in Streamflow on nutrient flux. Flux elasticities are less than unity (median = 0.93%) at most stations, but vary widely from 0.05% to 1.59%. Elasticities above unity occur most frequently in the largest rivers and in rivers draining the arid portions of the western Gulf of Mexico Basin. Historical flux variability and elasticity generally increase with the extent of arid conditions and the quantity of nonurban land use in the watershed. We extend the analysis of flux variability to examine several case studies of highly unusual meteorological events capable of significantly elevating nitrate flux and degrading estuarine ecology.