Carbon dioxide and submersed macrophytes in lakes: linking functional ecology to community composition.

Evaluating plant community response to atmospheric CO2 rise is critical to predicting ecosystem level change. Freshwater lakes offer a model system for examining CO2 effects as submersed macrophyte species differ greatly in their growth responses to CO2 enrichment, and free CO2 concentrations among these habitats show a wide range of natural, spatial variation. We determined free CO2 concentrations in the water column and sediment porewater in littoral zones with pH < 6.0 in Adirondack Mountain (New York, USA) lakes, and derived a community CO2 responsiveness index (CCRI) based on quantitative sampling of 15 submersed macrophyte communities coupled with greenhouse-derived growth responses to CO2 enrichment of constituent species to test two hypotheses: (1) CCRI, which is higher for communities dominated by species with greater growth responses to CO2 enrichment, is positively correlated to free [CO2 ] in the water column, and (2) in natural communities, the percent of sediment CO2 -using species, which are relatively unresponsive to CO2 enrichment, is negatively correlated to free [CO2 ]. A significant positive correlation (P = 0.003) between our physiologically based CCRI and the concentration of free CO2 in the water column supported our primary hypothesis that sites with higher levels of free CO2 are dominated by species with greater growth responses to CO2 enrichment. Our CCRI is also highly significantly correlated (P < 0.001) to the first axis scores for the same vegetation data from polar ordination. Finally, the relative importance of species that use sediment CO2 as a photosynthetic carbon source is significantly negatively correlated (P = 0.029) with the concentration of free CO2 in the water column. Our results indicate that natural variations in CO2 levels are important determinants of submersed macrophyte community composition. Further, we demonstrate the utility of a physiologically-based index of community composition, our CCRI, as an ecologically valid measure of community response to CO2 .

Positive feedback favors invasion by a submersed freshwater plant.

The submersed macrophyte Utricularia inflata has invaded lakes in northern New York State, thereby threatening native isoetids such as Eriocaulon aquaticum. Isoetids often dominate and modify softwater lakes due to their capacity to oxidize sediment and thus influence solute mobilization. Greenhouse experiments tested the hypotheses that U. inflata invasion could result in higher porewater iron (Fe) concentrations and greater ammonium (NH4 (+)) and Fe release from the sediment into the water column, and that this mobilization would stimulate further U. inflata growth. In the first experiment, three levels of U. inflata impact on E. aquaticum were imposed using sediment cores overlain by lake water: E. aquaticum alone, E. aquaticum with a cover of U. inflata, and bare sediment--the latter to simulate local extirpation of the isoetid by the invasive. After 16 weeks, sediment porewater NH4 (+) and total dissolved Fe concentrations were significantly higher (P < 0.05) for the U. inflata and bare sediment treatments. Water column concentrations of these solutes were five-fold higher (P < 0.05) for the bare sediment treatment than E. aquaticum alone, indicating that isoetid extirpation by U. inflata can compromise water quality. A second experiment demonstrated that U. inflata grew faster over bare sediment than over sediment with E. aquaticum (P < 0.05), likely due to greater solute mobilization in the absence of E. aquaticum. Where U. inflata causes a decline of native isoetids in Adirondack Mountain lakes, changes to lake sediment and water chemistry can create a positive feedback loop further escalating the impact of this invasive species.

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.

Physical site characteristics limit pollination and fruit set in the dioecious hydrophilous species, Vallisneria americana.

Experimental and observational studies of the submersed, freshwater macrophyte Vallisneria americana Michx. revealed that depth, wind and wave exposure, and current velocity may all influence fruit set. In this dioecious species, long-pedunculate female flowers are pollinated by free-floating male flowers at the water surface. Average fruit set in the natural populations studied varied from zero to 97% of the flowers observed. With increasing water depth in New York and Pennsylvania lakes, female plants continued to flower, though these flowers were unable to reach the surface, and consequently, did not set fruit. Fruit set was also lower in relatively open sites exposed to wind and waves, presumably because male flowers do not remain in the vicinity of female flowers long enough for effective pollination. This was particularly striking at a site with low male flower densities, but fruit set was increased to 100% at that site by confining the floating male flowers within a field enclosure. In a river, fruit set was negatively correlated with surface current velocity, and was reduced to zero in current velocities greater than 0.30 m · s-1. Fruit set in V. americana appears to be restricted or precluded by physical environmental conditions in a variety of sites.

Submersed macrophyte growth at low pH : II. CO2 × sediment interactions.

The submersed macrophyte Vallisneria americana was grown for seven weeks in a greenhouse to test for differences in the ability of three different sediments to support growth stimulation in response to CO2 enrichment at low pH. Plants accumulated 21- to 24-fold greater biomass at 10 × ambient CO2 concentrations than at ambient CO2 on all sediments. At both CO2 levels, plants grown on sediment from an acidified lake accumulated ca. 81%, and those grown on oligotrophic lake sediment ca. 47% as much biomass as plants grown on alkaline lake sediment. Despite striking CO2 and sediment effects on biomass accumulation, there was no significant interaction (using log-transformed data) between CO2 and sediment effects, indicating that all sediments allowed similar proportionate growth responses to CO2 enrichment. Plants grown on the less fertile sediments showed greater relative allocation to horizontal versus vertical growth by producing more rosette-bearing stolons in relation to plant height (leaf length) than plants grown on relatively fertile, alkaline lake sediment. Tissue analysis suggested that sediment effects on Vallisneria growth could be attributed neither to mineral putrient (nitrogen and phosphorus) limitation nor to aluminum toxicity in these low pH treatments. In any case, CO2 availability can be an important regulator of submersed macrophyte growth at low pH on a variety of sediment types, including those from oligotrophic and acidic lakes.

Submersed macrophyte growth at low pH : I. CO2 enrichment effects with fertile sediment.

Vallisneria americana was grown for six weeks in a greenhouse on relatively fertile sediment to test for factors other than nutrient limitation which may slow growth of this submersed macrophyte at pH 5. On the basis of dry mass accumulated, (1) low pH significantly depressed Vallisneria growth at constant free CO2 levels; (2) free CO2 enrichment, however, greatly stimulated Vallisneria growth at pH 5, by 2.8-fold and 10-fold at 3.2 times and 10 times air-equilibrated CO2 levels, respectively; and (3) growth was greater by far at pH 5 than at higher pH with constant total dissolved inorganic carbon (DIC). Free CO2 availability was thus an important controller of growth at low pH by Vallisneria americana on fertile sediment, and low pH was not directly deleterious. Field surveys of acidic lakes in the Adirondack Mountains of New York state revealed that DIC levels in low pH lakes were often well above equilibrium values and could potentially support vigorous macrophyte growth. Aluminum and/or iron toxicity did not appear to impair growth at low pH, and aluminum concentrations in Vallisneria shoots significantly decreased with increasing free CO2 concentrations at pH 5.0, perhaps due to growth dilution. Rosette production (a measure of asexual reproduction), maximum leaf length, and extent of flowering within treatments were positively correlated with plant biomass, rather than with pH or free CO2 levels per se.

Comparative desiccation tolerance of two Sphagnum mosses.

Sphagnum fallax (Klinggr.) Klinggr., a moss growing in hollows close to the water table, is more desiccation tolerant than S. nemoreum Scop., a hummock former distributed high above the hollows. Sphagnum fallax recovered to a greater proportion of its predesiccation photosynthetic rate after one and five days of tissue dryness. Further, a greater percentage of S. fallax plants survived five and ten day periods at low tissue water contents. Longer desiccated periods and lower water contents during these periods decreased both photosynthetic recovery and survival.Water contents measured in Bloomingdale Bog (Adirondack Mountains, NY, USA) showed that S. fallax probably dries more frequently and for longer periods than S. nemoreum. These results support previous findings that the greater ability of S. nemoreum to remain moist in the field is the most important character in its success as a hummock former. Greater tolerance of desiccation helps S. fallax to compensate for its greater tendency to become dry, and is a key physiological feature enabling it to dominate hollows.

Neighbor influences and seasonal growth patterns for Vallisneria americana in a mesotrophic lake.

Biomass of the submersed macrophyte Vallisneria americana rose steadily from early June to 50 g dry wt·m-2 in ealry August. Leaves dominated the biomass of this perennial from mid-June through August, after which plants senesced to leave only winter buds as perennating organs. Only 24% of plants sampled flowered during the 1978 growing season, yielding a population mean of less than 0.6% of dry weight allocated to sexual reproduction, with a maximum value of 7.8% for one plant. In contrast, allocation to winter buds accounted for 11% of total dry weight at the close of the growing season.Total dry weight, leaf length, stolon length, rosette number, and winter bud production of Vallisneria americana at the close of the growing season had significantly higher mean values for individual plants in a monospecific stand of Vallisneria than at sites dominated by Potamogeton amplifolius or Chara vulgaris. The basis of site differences in growth by V. americana is unknown, although we demonstrate significantly higher interstitial soluble reactive phosphorus concentration and organic content of sediment at a Vallisneria site.Removal of neighbors of some Vallisneria plants at the Vallisneria, Potamogeton, and Chara sites resulted in longer leaves, but fewer rosettes for plants with neighbors than plants without neighbors. It appears that this changing growth pattern permitted Vallisneria plants to compensate for the presence of neighbors, and thereby attain similar biomass and sustain similar winter bud production in the presence and absence of neighbors.

Coexistence and the comparative light relations of the submersed macrophytes Myriophyllum spicatum L. and Vallisneria americana Michx.

The Eurasian watermilfoil (Myriophyllum spicatum L.) has partially replaced wild celery (Vallisneria americana Michx.) as a community dominant in the littoral zones of lakes of Madison, Wisconsin. The two species have very different growth forms, with that of M. spicatum corresponding more closely to the optimal growth form simulated by the macrophyte production model WEED. The objective of this research was to investigate the mechanisms by which Vallisneria could compensate for its nonoptimal growth form and coexist with Myriophyllum.A quantification of midsummer growth form for the two species at a rooting depth of 80-90 cm showed that M. spicatum had 68% of its shoot biomass within 30 cm of the surface, whereas V. americana had 62% of its leaf biomass within 30 cm of the bottom. Vallisneria had a light extinction coefficient ranging from 0.013 to 0.019 m2·g-1, much higher than the value (ca. 0.006 m2·g-1) for M. spicatum. This indicates less effective penetration of light to lower leaves of V. americana. Half-saturation constants describing the light-dependence of carbon uptake in "shade" and "sun" tissues ranged from 60-197 microeinsteins·m-2·s-1 for V. americana, and 164-365 µeinsteins·m-2·s-1 for M. spicatum. The optimum temperature for photosynthesis was 33.6°C for M. spicatum and 32.6°C for V. americana, but Myriophyllum was nearly twice as effective at carbon uptake at 10°C. Integration of all of the above features with WEED showed that, for midsummer conditions, V. americana more than compensated for apparently disadvantageous morphological features by its greater physiological adaptability to low light regimes. Coupled with the temperature-dependence of photosynthesis, it appears that V. americana is favored by midsummer conditions, whereas M. spicatum is at an advantage at other times.