Temporal and spatial variation in flower and fruit production in a food-deceptive orchid: a five-year study

Deceptive orchids are generally characterized by low levels of fruit set; however, there may be substantial variations in fruit set between sites and years. Within a single population, individual plants may also differ greatly in their reproductive output as a result of differences in inflorescence size or local density. In this study, we determined flower and fruit production over 5 years in two populations of the food-deceptive orchid, Orchis purpurea . All plants were monitored annually for survival and flowering at each site to determine whether flowering and fruiting induced costs. The number of flowers per inflorescence varied considerably from year to year (min: 36.6, max: 49.5). Average fruit set was low (7%) and varied considerably among years and populations. A considerable proportion of plants also failed to set any fruit. However, the probability of producing at least one fruit was not affected by inflorescence size or local density. The number of fruits was significantly related to inflorescence size, but proportional fruit set was not. Local density also did not affect the number of fruits, nor proportional fruit set. There was also no evidence that plants with large inflorescence size or high fruiting success had a larger probability of remaining vegetative the year after flowering than plants with small inflorescence size or low fruiting success. Our results suggest that pollinator-mediated selective forces on inflorescence size through female reproductive success alone are weak, most likely because of the low overall level of visitation and the resulting uncertainty of pollination at the individual level. Our results further demonstrate that investigation of patterns of fruit set over several years is needed to better understand the variability in female reproductive success that is typical of most plant–pollinator interactions.     

Strong growth limitation of a floating plant (Lemna gibba) by the submerged macrophyte (Elodea nuttallii) under laboratory conditions

1. The asymmetric competition for light and nutrients between floating and submerged aquatic plants is thought to be key in explaining why dominance by either of these groups can be stable and difficult to change. 2. Although the shading effect of floating plants on submerged plants has been well documented, the impact of submerged plants on floating plants has been poorly explored hitherto. 3. Here, we used laboratory experiments to examine how submerged plant (Elodea nuttallii) alter nutrient conditions in the water column and how this affects the growth of floating plants (Lemna gibba). 4. We demonstrate that, at higher nutrient concentrations, Lemna is increasingly likely to outcompete Elodea. 5. Under low nutrient concentrations (0.1–2 mg N L^?1) Elodea can strongly reduce the growth of Lemna. Growth of floating plants virtually stopped in some of the experiments with Elodea. 6. Extremely reduced tissue N, Mn, chlorophyll and elongated roots indicated that the growth inhibition of Lemna by Elodea was predominantly caused by the latter’s impact on the nutrient conditions for floating plants. 7. These results strengthen the hypothesis that submerged plants can prevent colonization of a lake by floating plants.     

Microbial enzyme activity,nutrient uptake and nutrient limitation in forested streams

1. We measured NH₄⁺ and PO₄^?3 uptake length (S_w), uptake velocity (V_f), uptake rate (U), biofilm respiration and enzyme activity and channel geomorphology in streams draining forested catchments in the northwestern (Northern California Coast Range and Cascade Mountains) and southeastern (Appalachian and Ouachita mountains) regions of the United States. Our goal was to use measures of biofilm enzyme activity and nutrient uptake to assess nutrient limitation in forested streams across broad regional scales. 2. Geomorphological attributes, biofilm enzyme activity and NH₄⁺ uptake were significantly different among streams in the four study units. There was no study unit effect on PO₄^?3 uptake. The proportion of the stream channel in pools, % woody debris, % canopy closure, median substrate size (d₅₀), stream width (w), stream velocity (v), discharge (Q), dispersion coefficient (D) and transient storage (A_s/A) were correlated with biofilm enzyme activity and nutrient uptake in some study units. 3. Canonical correlation analyses across study units revealed significant correlations of NH₄‐V_f and PO₄‐V_f with geomorphological attributes (w, d₅₀, D, % woody debris, channel slope and % pools) and biofilm phosphatase activity. 4. The results did not support our expectation that carbon processing rates by biofilm microbial assemblages would be governed by stream nutrient availability or that resulting biofilm enzyme activity would be an indicator of nutrient uptake. However, the relative abundances of peptidases, phosphatase and glycosidases did yield insight into potential N‐, P‐ and C‐limitation of stream biofilm assemblages, and our use of biofilm enzyme activity represents a novel application for understanding nutrient limitations in forested streams. 5. Regressions of V_f and U against ambient NH₄⁺ and PO₄^?3 indicated that none of our study streams was either NH₄⁺ or PO₄^?3 saturated. The Appalachian, Ouachita and Coastal streams showed evidence of NH₄⁺ limitation; the Ouachita and Coastal streams were PO₄^?3 limited. As a correlate of nutrient limitation and saturation in streams, ratios of total aminopeptidase and phosphatase activities and the ratio of NH₄‐U to PO₄‐U indicate these forested streams are predominantly N‐limited, with only the streams draining Ouachita and Coastal catchments demonstrating appreciable levels of P‐limitation. 6. Our results comparing the stoichiometry of microbial enzyme activity with nutrient uptake ratios and with the molar ratios N and P in stream waters suggest that biological limitations are not strictly the result of stream chemistry and that the assessments of nutrient limitations in stream ecosystems should not be based on chemistry alone. 7. Our present study, along with previous work in streams, rivers and wetlands, suggests that microbial enzyme activities, especially the ratios of total peptidases to phosphatase, are useful indicators of nutrient limitations in aquatic ecosystems.     

Interpretation of metabolic flux maps by limitation potentials and constrained limitation sensitivities

Two new concepts, "Limitation Potential" and "Constraint Limitation Sensitivity" are introduced that use definitions derived from metabolic flux analysis (MFA) and metabolic network analysis (MNA). They are applied to interpret a measured flux distribution in the context of all possible flux distributions and thus combine MFA with MNA. The proposed measures are used to quantify and compare the influence of intracellular fluxes on the production yield. The methods are purely based on the stoichiometry of the network and constraints that are given from irreversible fluxes. In contrast to metabolic control analysis (MCA), within this approach no information about the kinetic mechanisms are needed. A limitation potential (LP) is defined as the reduction of the reachable (theoretical) maximum by a measured flux. Measured fluxes that strongly narrow the reachable maximum are assumed to be limiting as the network has no ability to counterbalance the restriction due to the observed flux. In a second step, the sensitivity of the reduced maximum is regarded. This measure provides information about the necessitated changes to reach higher yields. The methods are applied to interpret the capabilities of a network based on measured fluxes for a L-phenylalanine producer. The strain was examined by a series of experiments and three flux maps of the production phase are analyzed. It can be shown that the reachable yield is drastically reduced by the measured efflux into the TCA cycle, while the oxidative pentose-phosphate pathway only plays a secondary role on the reachable maximum.     

Importance of sediment deposition and denitrification for nutrient retention in floodplain wetlands

Questions: Various floodplain communities may differ in their relative abilities to influence water quality through nutrient retention and denitrification. Our main questions were: (1) what is the importance of sediment deposition and denitrification for plant productivity and nutrient retention in floodplains; (2) will rehabilitation of natural floodplain communities (semi‐natural grassland, reedbed, woodland, pond) from agricultural grassland affect nutrient retention? Location: Floodplains of two Rhine distributaries (rivers Ussel and Waal), The Netherlands. Methods: Net sedimentation was measured using mats, denitrification in soil cores by acetylene inhibition and bio‐mass production by clipping above‐ground vegetation in winter and summer. Results: Sediment deposition was a major source of N and P in all floodplain communities. Highest deposition rates were found where water velocity was reduced by vegetation structure (reedbeds) or by a drop in surface elevation (pond). Sediment deposition was not higher in woodlands than in grassland types. Denitrification rates were low in winter but significantly higher in summer. Highest denitrification rates were found in an agricultural grassland (winter and summer) and in the ponds (summer). Plant productivity and nutrient uptake were high in reedbeds, intermediate in agricultural grasslands, ponds and semi‐natural grasslands and very low in woodlands (only understorey). All wetlands were N‐limited, which could be explained by low N:P ratios in sediment. Conclusions: Considering Rhine water quality: only substantial P‐retention is expected because, relative to the annual nutrient loads in the river, the floodplains are important sinks for P, but much less for N. Rehabilitation of agricultural grasslands into ponds or reedbeds will probably be more beneficial for downstream water quality (lower P‐concentrations) than into woodlands or semi‐natural grasslands.     

Coupled Nitrogen and Calcium Cycles in Forests of the Oregon Coast Range

Nitrogen (N) is a critical limiting nutrient that regulates plant productivity and the cycling of other essential elements in forests. We measured foliar and soil nutrients in 22 young Douglas-fir stands in the Oregon Coast Range to examine patterns of nutrient availability across a gradient of N-poor to N-rich soils. N in surface mineral soil ranged from 0.15 to 1.05% N, and was positively related to a doubling of foliar N across sites. Foliar N in half of the sites exceeded 1.4% N, which is considered above the threshold of N-limitation in coastal Oregon Douglas-fir. Available nitrate increased five-fold across this gradient, whereas exchangeable magnesium (Mg) and calcium (Ca) in soils declined, suggesting that nitrate leaching influences base cation availability more than soil parent material across our sites. Natural abundance strontium isotopes (⁸⁷Sr/⁸⁶Sr) of a single site indicated that 97% of available base cations can originate from atmospheric inputs of marine aerosols, with negligible contributions from weathering. Low annual inputs of Ca relative to Douglas-fir growth requirements may explain why foliar Ca concentrations are highly sensitive to variations in soil Ca across our sites. Natural abundance calcium isotopes (δ⁴⁴Ca) in exchangeable and acid leachable pools of surface soil measured at a single site showed 1 per mil depletion relative to deep soil, suggesting strong Ca recycling to meet tree demands. Overall, the biogeochemical response of these Douglas-fir forests to gradients in soil N is similar to changes associated with chronic N deposition in more polluted temperate regions, and raises the possibility that Ca may be deficient on excessively N-rich sites. We conclude that wide gradients in soil N can drive non-linear changes in base-cation biogeochemistry, particularly as forests cross a threshold from N-limitation to N-saturation. The most acute changes may occur in forests where base cations are derived principally from atmospheric inputs.     

Patterns in potassium dynamics in forest ecosystems

The biotic cycling of potassium (K) in forest systems has been relatively understudied in comparison with nitrogen (N) and phosphorus (P) despite its critical roles in maintaining the nutrition of primary production in forests. We investigated the ecological significance of K in forests from a literature review and data synthesis. We focused on (1) describing patterns of the effects of K availability on aboveground growth and change in foliar tissue of tree species from a variety of forests; and (2) documenting previously unreported relationships between hydrologic losses of K and N in forested watersheds from the Americas. In a review of studies examining tree growth under K manipulations/fertilizations, a high percentage (69% of studies) showed a positive response to increases in K availability in forest soils. In addition, 76% of the tree studies reviewed showed a positive and significant increase in K concentrations in plant tissue after soil K manipulation/fertilization. A meta-analysis on a subset of the reviewed studies was found to provide further evidence that potassium effects tree growth and increased tissue [K] with an effect size of 0.709 for growth and an overall effect size of 0.56. In our review of watershed studies, we observed that concentrations of K typically decreased during growing seasons in streams draining forested areas in the Temperate Zones and were responsive to vegetation disturbance in both temperate and tropical regions. We found a strong relationship (r2 = 0.42-0.99) between concentrations of K and N (another critical plant nutrient) in stream water, suggesting that similar mechanisms of biotic retention may control the flow of these nutrients. Furthermore, K dynamics appear to be unique among the base cations, e.g. calcium, magnesium, and sodium, because the others do not show similar seasonal patterns to K. We suggest that K may be important to the productivity and sustenance of many forests, and its dynamics and ecological significance warrant further study. We suggest that knowledge about the dynamics of this understudied element is imperative for our understanding patterns and processes in forest ecosystems.     

Iron limits primary productivity during spring bloom development in the central North Atlantic

We present in situ biophysical measurements and bioassay experiments that demonstrate iron limitation of primary productivity during the spring bloom in the central North Atlantic. Mass balance calculations indicate that nitrate drawdown is iron (Fe)-limited and that aeolian Fe supply to this region cannot support maximal phytoplankton growth during the bloom. Using a simple simulation model, we show that relief of Fe limitation during the spring bloom can increase nitrate drawdown and, hence, new primary production, by 70%. We conclude that the episodic nature of iron supplied by dust deposition is an important factor controlling the dynamics of the spring bloom. From this, we hypothesize that variability in the timing and magnitude of the spring bloom in response to aeolian Fe supply will affect carbon drawdown and food web dynamics in the central North Atlantic.