Phosphorus fluctuation in water and deposition into sediment within an emergent macrophyte stand

Seasonal fluctuation of phosphorus in water, and total phosphorus and different inorganic P fractions (extracted by NH₄Cl, NH₄F, NaOH and H₂SO₄) and organic P fraction (residual P) in surface sediment, were measured in the littoral of oligotrophic Lake Pääjärvi (southern Finland). After the emergence of Equisetum fluviatile L. shoots in mid June, water exchange between the littoral and pelagial area diminished and phosphorus concentrations in water and in surface sediment increased in the inner and mid littoral zones. Phosphorus pool in flocculent, easily resuspensible sediment composed on average 62% of inorganic phosphorus and 38% of organic phosphorus. 63% of the inorganic phosphorus, on the average, was extracted from apatite-P fraction 29% from iron-P fraction 7% from aluminum-P fraction and less than 1% from loosely-bound P fraction. During the growing season, net accumulation of sediment and phosphorus was measured only in the inner littoral zone whereas the outer littoral zones acted rather as a source of phosphorus for the pelagic area. The results of this study indicated that nutrient dynamics in littoral environment was through changes in flow environment greatly governed by the macrophytes.     

Shading by an invasive macrophyte has cascading effects on sediment chemistry

The submersed freshwater macrophyte Utricularia inflata is a recent invader of Adirondack Mountain lakes (NY, USA). Previous experiments suggested that U. inflata can indirectly change nutrient cycling in Adirondack lake ecosystems by reducing the growth of native isoetid macrophytes, which in turn affects sediment chemistry. A 13-week greenhouse experiment was conducted to test the hypothesis that shading can explain the detrimental effect of U. inflata on the native short-statured isoetid, Eriocaulon aquaticum. Eriocaulon aquaticum has a dense root system that oxidizes sediment by releasing oxygen; it also takes up carbon dioxide from sediment. Growth and asexual reproduction of E. aquaticum grown under shaded conditions was reduced significantly compared to an unshaded control (P < 0.001). Shading resulted in sediment changes: redox potential fell from 216 mV in the absence of shading to 76 mV under four layers of shade cloth (P < 0.0001). Shading also increased the concentration of extractable sediment ammonium (P < 0.01), as well as carbon dioxide concentrations (P < 0.0001) and pH of porewater (P < 0.05). The effect of U. inflata on the native isoetids and consequently on sediment chemistry closely matched the impact of shade cloth with similar light attenuation. Our results indicate that the principal mechanism by which U. inflata affects native isoetids and sediment chemistry is shading.     

Growth and morphological responses to water level and nutrient supply in three emergent macrophyte species

Sanjiang Plain is the largest freshwater marsh in China, where plant zonation along water-level gradients is a common phenomenon. The aim of this experiment was to identify the role of water level and nutrient availability on plant zonation in the plain. Growth and root morphology of three perennial emergent macrophyte species were investigated by growing in two water levels (0.1 and 10.0 cm, relative to soil surface) and in two levels of nutrient supply (0 and 0.5 g slow-release fertilizer per container). In the plain, Carex lasiocarpa typically occurs at low elevations, Glyceria spiculosa at medial elevations, and Deyeuxia angustifolia at high elevations. The relative growth rate was the highest in C. lasiocarpa and the lowest in D. angustifolia in the 10.0-cm water level. Among the three species, only total biomass of D. angustifolia was affected by water level, and decreased with increasing water level. High nutrient supply led to increased total biomass in C. lasiocarpa and G. spiculosa. High water level led to an increased root diameter in G. spiculosa and a decreased root length in C. lasiocarpa. In the 10.0-cm water level, low nutrient supply led to thinner roots in D. angustifolia, but resulted in an increased specific root length (SRL) in C. lasiocarpa and root diameter in G. spiculosa. Water-level effect on root porosity was only observed in G. spiculosa, and nutrient amendment did not influence root porosity in all the species. These data indicate that both nutrient and water level are important factors regulating plant distribution pattern in the Sanjiang Plain, because both C. lasiocarpa and G. spiculosa are relatively sensitive to nutrient supply whereas D. angustifolia is sensitive to water level. Handling editor: S. M. Thomaz     

Transpiration/biomass ratio for carrots as affected by salinity,nutrient supply and soil aeration

The influence of salinity, nutrient level and soil aeration on the transpiration coefficient, defined as amount of water transpired/unit biomass produced (transpiration/biomass ratio) of carrots was investigated under non-limiting conditions with respect to water supply.Under optimum conditions and favorable nutrient supply, the transpiration coefficient amounted to 280–310 g H₂O g^–1 storage root dry weight (RDW). The transpiration coefficient did not change significantly up to salt concentration of 16 mS cm^–1 in the soil solution under otherwise optimum conditions. Higher salt concentrations or low nutrient levels increased the transpiration coefficient to values of 390–540 g H₂O g^–1 RDW. It is suggested that the transpiration coefficient is not affected by salinity as long as toxic effects and nutrient imbalances do not occur. The transpiration coefficient was not increased by impeded soil aeration. Biomass production was more negatively influenced by adverse soil conditions (salinity, low nutrient level, impeded soil aeration) than was the transpiration coefficient.     

Growth of the seaweed Ulva rigida C. Agardh in relation to biomass densities, internal nutrient pools and external nutrient supply in the Sacca di Goro lagoon (Northern Italy)

Growth of the seaweed Ulva rigida C. Agardh was investigated in relation to biomass densities, internal nutrient pools and external nutrient supply. Research was carried out from 23 March to 5 July 1994 in the Sacca di Goro (Po Delta, Northern Italy), whose south-eastern part was covered by extensive mats of Ulva rigida. Two types of field experiments were conducted by incubating Ulva thalli inside large cages. In the first experiment, beginning on 23 March, 100 g of wet thalli were placed into the cages, allowed to grow for two weeks, then collected and replaced. This procedure was repeated 8 times over the study period. In the second experiment, Ulva thalli were left inside the cages and collected at selected time intervals (14, 27, 41, 64 and 76 days) in order to simulate the effects of increased density on growth and nutrient storage.We recorded specific growth rates (NGR) ranging from 0.025 to 0.081 d^–1 for a period up to two months in the repeated short-term experiments performed at relatively low initial algal densities (300–500 g AFDW m^–3). These NGR resulted significantly related to dissolved inorganic nitrogen (DIN) in the water column. Tissue concentrations of total Kjeldahl nitrogen (TN) were almost constant, while extractable nitrate decreased in a similar manner to DIN in the water column. Total phosphorus showed considerable variation, probably linked to pulsed freshwater inflow.In the long-term incubation experiment, NGR of Ulva was inversely related to density. Internal concentrations of both total P and TN reached maximum values after one month; thereafter P concentration remained almost constant, while TN decreased below 2% w/w (by dry weight). The TN decrease was also accompanied by an abrupt decrease in nitrate tissue concentration. The biomass incubated over the two month period suffered a progressive N limitation as shown by a decreasing NY ratio (49.4 to 14.6). The reciprocal control of Ulva against biogeochemical environment and viceversa is a key factor in explaining both resource competition and successional stages in primary producer communities dominated by Ulva. However, when the biomass exceeds a critical threshold level, approximately 1 kg AFDW m^–3, the macroalgal community switches from active production to rapid decomposition, probably as a result of selfshading, biomass density and development of anaerobic conditions within the macroalgal beds.     

Transpiration and root water uptake by olive trees

While the cultivated olive tree (Olea europaea L.) is known to be sclerophyllous and effective at tolerating drought, little is known of its short-term water-use dynamics for most studies have been based on longer-term, water-balance information. We present here, for the first time, heat-pulse measurements of the sap flux measured not only within the semi-trunk of an olive tree, but also within a root excavated close to the stump. One tree in the olive grove near Seville in Spain had regularly received basin irrigation during the summer, whereas the other, growing on this deep silt loam, had been without water for over 3 months. Following a flood irrigation of 730 L to a dyked area around the tree, the regularly-irrigated olive maintained a transpiration rate of 1.65 mm³ mm^–2 d^–1, on a leaf area basis, for only 3 days following the irrigation. This rate was maintained for a total consumption of 110 L. It then began again to limit its rate of water use with transpiration falling below that predicted for well-watered conditions by the Penman-Monteith equation. The flow of sap in the near-surface root dropped concomitantly. Meanwhile the unirrigated tree was using water at just 0.78 mm d^–1. Yet following an irrigation of 870 L it only lifted its consumption to 1.12 mm d^–1, on a leaf area basis. Neither did it recover its leaf water potential following this wetting because of an inability to refill cavitated vessels. These data again show olive to be a parsimonious and cautious consumer of soil water.     

Growth strategy of an emergent macrophyte, Typha orientalis Presl, in comparison with Typha latifolia L. and Typha angustifolia L.

The growth strategy of an emergent plant, Typha orientalis Presl, was examined in experimental ponds in comparison with two other Typha species distributed in Japan, Typha latifolia L. and Typha angustifolia L. T. orientalis showed the greatest ability of vegetative reproduction at the expense of growth in height. T. orientalis started to produce new ramets earlier than T. latifolia and T. angustifolia. These results suggest that T. orientalis should be a rather pioneer-like species and would be restricted to disturbed habitats.     

An invasive macrophyte alters sediment chemistry due to suppression of a native isoetid

The submersed macrophyte Utricularia inflata (inflated bladderwort) is a recent invader of Adirondack Mountain lakes (NY, USA). A 15-week greenhouse experiment and a 7-week field experiment were conducted to test the hypothesis that this rootless species fundamentally changes sediment chemistry through its suppression of the native short-statured species, Eriocaulon aquaticum. E. aquaticum has an extensive root system that releases oxygen into the sediment. In greenhouse conditions, E. aquaticum raised the porewater redox potential of otherwise bare sediment from 25 to 324 mV, lowered the sediment porewater pH from 5.7 to 4.6, and depleted the dissolved inorganic carbon and ammonium concentrations in the sediment porewater by 68.4 and 96.0%, respectively (P<0.001 for all four parameters). A cover of U. inflata over E. aquaticum, however, greatly reduced the latter’s effect on redox potential (P<0.001), dissolved solutes (P<0.001), and pH (P<0.05). E. aquaticum biomass increased during the greenhouse experiment in the absence of U. inflata, but decreased in its presence (P<0.001). Redox and growth rate results from the field experiment paralleled those from the greenhouse experiment. Our data suggest that U. inflata may change nutrient cycling in Adirondack lake ecosystems by reducing the growth of native isoetid macrophytes, such as E. aquaticum, and consequently altering key features of sediment chemistry.     

Physiological integration helps a clonal macrophyte spread into competitive environments and coexist with other species

Physiological integration may help clonal macrophytes invade or escape from existing communities. No studies have tested the above hypothesis in aquatic plants. In an outdoor pond experiment, we subjected clonal fragments of the submerged macrophyte Vallisneria spiralis L. to heterogeneous environments in which V. spiralis spread from bare habitats towards vegetated habitats occupied by Myriophyllum spicatum L. or V. spiralis spread from vegetated habitats towards bare habitats. V. spiralis stolons between ramets in bare habitats and in vegetated habitats were either intact or severed. We investigated the habitat selection of V. spiralis by examining the allocation of biomass and ramets to heterogeneous habitats during its vegetative spread phase. Results showed that the stolon connection had different effects on the habitat selection of V. spiralis with regard to invasion and escape. When V. spiralis spread from bare to vegetated habitats, in comparison to severing the stolon, the stolon connection eventually facilitated a 49% increase in biomass and a 27% increase in number of ramets allocated to vegetated habitats. However, when V. spiralis spread from vegetated to bare habitats, biomass and ramets allocated to bare habitats were not significantly changed by the stolon connection (only a 5% increase in biomass and a 6% increase in number of ramets). These results indicate that clonal integration facilitated V. spiralis not to escape from but invade into vegetated habitats. The study provides evidence that physiological integration is important for survival and tolerance of ramets in competitively stressful environments and can help clonal macrophytes coexist with other species.     

The water velocity preferences of Austrosimulium furiosum and Simulium ornatipes (Diptera: Simuliidae) larvae,and the implications for micro-habitat partitioning

The aquatic larvae of two simuliid species, Austrosimulium furiosum (Skuse) and Simulium ornatipes Skuse, which often occur together in Victorian streams, were shown in laboratory experiments to have preferences for different water velocities: larvae of A. furiosum preferred water velocities of 0.2–0.3 m s^–1, and S. ornatipes preferred water velocities of 0.9–1.3 m s ^–1 . Final instar larvae of both species selected slow water speeds of less than 0.25 m s ^–1 prior to pupation. Flow patterns around a cylinder in a laboratory stream were mapped, and the distribution of A. furiosum larvae within the wake, paired vortices and horseshoe vortex was recorded. Larvae selected areas with suitable water velocities and aligned with the flow, providing flow visualization of micro-current speed and direction. The potential for micro-habitat partitioning is discussed in relation to benthic sampling strategies.     

Effects of water and nutrient supply on amount and on nutrient concentration of litterfall and forest floor litter in Eucalyptus globulus plantations

A field experiment was initiated in March 1986 in central Portugal to evaluate the influence of water and nutrient supply on the productivity of E. globulus. The treatments applied were, irrigation plus fertiliser, irrigation and the application of fertilisers in rainfed plots. The control received neither fertilisers nor irrigation. The annual pattern litterfall was measured over a period of 5 years and the litter layer was quantified 6 years after planting. The amount of litterfall varied with the treatments. Simultaneous water and nutrient supply increased significantly litterfall, in respect to control. In rainfed conditions the timing of the maximum of litterfall was anticipated relative to the irrigated plots. However, the time of maximum litterfall did not coincide with the dry season but rather with period of maximum growth in each treatment. The N and P concentration was higher in the litterfall of the two fertilised treatments than in the others. The lowest concentrations of N and P in the leaf litter were coincident with the summer period in all treatments. The withdrawal of N and P ranged between 32 and 65% according to treatment and season. The deliverance of nutrients through litterfall was strongly increased by simultaneous water and nutrient supply. The supply of fertilisers in rainfed conditions promoted higher deliverance of nutrients than in the plots irrigated only. The mass of the litter layer was significantly increased by simultaneous water and nutrient supply. Application of fertilisers induced an increase in N and P concentration and a decrease in C/N ratio of the litter layer. Treatments and C/N values did not influence apparently the proportion of carbon, N and P released through mineralization from the litter layer. The non-fertilized treatments showed a more efficient N cycling than the others.     

A comparison of nutrient availability measured by chemical analysis and calculated from bioassay yields

The concentrations of soluble biologically available nitrogen and phosphorus were determined using Selenastrum capricornutum bioassays and compared with analytically measured soluble nitrate (NO₃-N) and soluble reactive phosphorus concentrations during enrichment studies in a South African impoundment. The NO₃-N analyses consistently underestimated the soluble biologically available nitrogen and the extent of the discrepancy decreased with increasing NO₃-N concentration. Biological availability of soluble organic nitrogen during the bioassays is suggested as a reason for the discrepancies. At low soluble reactive phosphorus concentrations the analytical measurements underestimated the soluble biologically available phosphorus while at high soluble reactive phosphorus concentrations the analytical measurements were considerable overestimates of soluble biologically available phosphorus. Possible reasons for the observed trend are discussed.     

The effects of varying culture nitrogen and phosphorus levels on nutrient uptake and storage by the waterhyacinth Eichhornia crassipes (Mart) Solms

The uptake of nitrogen (N) by waterhyacinth (Eichhornia crassipes) was maximal when the culture solution contained a combination of 36 ppm N and 6,53 ppm phosphorus (P). N uptake was inhibited by increasing P or decreasing N in the culture medium. Uptake of P was stimulated by the elevation of either N or P or both. An increase of P above O ppm inhibited the accumulation of N. Similarly, the accumulation of P was enhanced by increasing levels of P but was reduced with increasing levels of N. Both N and P levels were always greatest in leaves; next highest in floats and lowest in roots. High tissue levels of these two nutrients and the capacity for nutrient removal from water by waterhyacinth indicates a possible role for this plant in the reduction of eutrophication in nutrient polluted waters as well as the use of harvested, dried tissue for fertilizer.     

Soil solution phosphate,root uptake kinetics and nutrient acquisition: implications for a patchy soil environment

Summary The importance of increased root phosphate (P) uptake kinetics, root proliferation and local increases of soil solution P (P₁) for P acquisition from fertile soil microsites was explored with a simulation model and calculated uptake was compared with experimental data. Based on the partitioning of added P in microsites to P₁ and P adsorbed on soil particles and the results of a dual-isotope-labeling experiment (Caldwell et al. 1991a), acquisition of P from the fertile microsites was some 20 X that of uptake from an equal volume of soil which received only water. Simulations were in general agreement and also showed that elevation of root P uptake kinetics could contribute more to the increased acquisition than did root proliferation under these circumstances. Although increased physiological uptake capacity for P has generally been considered to be of little benefit because of diffusion limitation, in patchy soil environments selective elevation of P uptake kinetics in fertile microsites may be of considerable benefit. These tests were conducted in calcareous soil which releases much less P into the soil solution than do many other soils. In many noncalcareous soils the benefits of selective elevation of root uptake kinetics would likely be greater.     

Isotopic enrichment in a phloem-feeding insect: influences of nutrient and water availability

The isotopic enrichment between an animal and its diet can vary among and within living systems, but the sources of variation are not yet fully understood. Some studies have found that diet quality or an animal’s nutritional status can influence the degree of trophic enrichment, while others have dismissed nutrition as a contributing factor. We evaluated the effects of nutrient and water availability on carbon and nitrogen isotopic enrichment in a specialized plant–herbivore system. Aphids are largely sedentary and rely exclusively on nitrogen-poor phloem sap of their host for nutrition. We grew potato aphids [Macrosiphum euphorbiae (Aphididae)] on an accepted host, pumpkin [Cucurbita pepo L. (Cucurbitaceae)], in a glasshouse environment. Twelve pumpkin plants growing under high- and low-watering regimes were inoculated at 4 weeks of age with aphids. During the course of the experiment we collected leaves, phloem sap, aphids and honeydew (i.e., aphid exudates). We found no trophic enrichment between aphids and their phloem sap diet, but significant carbon enrichment of honeydew relative to aphids (2.5‰) and phloem sap (2.1‰). Honeydew was also enriched in nitrogen compared to the phloem sap (1.2‰). Watering treatment had a substantial impact on trophic enrichment. Correlations among tissues, an indication of uniform trophic enrichment among samples, were significant only for the carbon isotopic composition, and then only for plants and aphids grown in the low-water treatment. Diet quality also influenced the degree of isotopic enrichment; trophic enrichment for both carbon and nitrogen isotopic composition increased as diet quality (C/N) declined. We conclude that the degree of trophic enrichment is variable due, in part, to diet quality, but that the scale of variation is small.     

Difference in the hypoxia tolerance of the round crucian carp and largemouth bass: implications for physiological refugia in the macrophyte zone

The hypoxia tolerance of larval and juvenile round crucian carp, Carassius auratus grandoculis, and largemouth bass, Micropterus salmoides, was determined using respirometry to examine the potential of hypoxic areas in the macrophyte zone as physiological refugia for round crucian carp. The tolerance, which was measured as the critical oxygen concentration (Pc), was 1.32 mg O₂/l in the round crucian carp and 1.93 mg O₂/l in the largemouth bass. As the round crucian carp tolerated hypoxia better than the largemouth bass, hypoxic areas in the macrophyte zone might function as physiological refugia for round crucian carp.     

Biomass and photosynthetic productivity of water hyacinth (Eichhornia crassipes) as affected by nutrient supply and mirid (Eccritotarus catarinensis) biocontrol

The biological control of water hyacinth is affected by water nitrogen and phosphorus content and this was investigated experimentally at five levels of nutrient supply by measuring plant photosynthetic and growth responses, and mirid reproduction and herbivory of nutrient treated plants. Low nitrogen (2–0.2 mg L⁻¹) and phosphorus (0.2–0.01 mg L⁻¹) supply decreased hyacinth photosynthesis, growth and biomass accumulation relative to plants supplied 200 mg L⁻¹ N and 20 mg L⁻¹ P. This effect depended more on nitrogen supply than phosphorus supply. Chlorophyll fluorescence showed that the photosynthetic light reactions of low nutrient plants were affected and leaves had decreased chlorophyll content, density of functional photosystems II and dissipated a greater proportion of absorbed energy as heat. Gas exchange parameters showed reduced carboxylation efficiency, rates of RuBP regeneration and light saturated photosynthetic rates, but not quantum yields. Effects on photosynthesis translated into lower plant dry biomass. Mirid herbivory exacerbated the effects of low nutrients noted for chlorophyll fluorescence, gas exchange parameters and biomass accumulation, however, these effects were not always significant and there was no obvious correlation between the level of nutrients supplied and the effect of mirid herbivory. Low nutrient supply did, however, affect mirid performance reducing the number of adult insects, nymphs and herbivory intensity suggesting that in the long-term mirid populations would be significantly affected by water nutrient status.     

The correlations between system treatment efficiencies and aboveground emergent macrophyte nutrient removal for the Hsin-Hai Bridge phase II constructed wetland

This study investigated the correlations between the system treatment efficiencies and total nitrogen (TN) and total phosphorus (TP) accumulations of aboveground tissues of the wetland macrophytes of Hsin-Hai Bridge phase II constructed wetland. Among 19 emergent macrophytes studied, the optimal TN contents, 3.82% and 3.52% (w/w) were found for water spinach (Ipomoea aquatica) and Ludwigia x taiwanensis; while the optimal TP contents were found for the above two macrophytes at 0.64% and 0.83% (w/w). The accumulations of total plant TN and TP uptakes increased from 213 to 403 kg and 41 to 75 kg from March 2007 to the peak at September 2007, respectively. The TN ratios between plant tissue accumulations and the removals from the influents were 1.57%, 2.76%, 1.51% and 3.2% from March 2007 to March 2008. In the same period, the TP ratios between plant tissue accumulations and the removals from influents were 1.71%, 8.0%, 0.58% and 10.1%. The roles of the uptakes by aboveground portions of emergent macrophytes in system nutrient removals from the influents were more significant during growth seasons.