INORGANIC CARBON TRANSPORT IN RELATION TO CULTURE AGE AND INORGANIC CARBON CONCENTRATION IN A HIGH-CALCIFYING STRAIN OF EMILIANIA HUXLEYI (PRYMNESIOPHYCEAE)1

The relationships among inorganic carbon transport, bicarbonate availability, intracellular pH, and culture age were investigated in high-calcifying cultures of Emiliania huxleyi (Lohmann) Hay & Mohler. Measurement of inorganic carbon transport by the silicone-oil centrifugation technique demonstrated that gadolinium, a potential Ca²⁺ channel inhibitor, blocked intracellular inorganic carbon uptake and photosynthetic ¹⁴CO₂₊ fixation in exponential-phase cells. In stationary-phase cells, the intracellular inorganic carbon concentration was unaffected by gadolinium. Gadolinium was also used to investigate the link between bicarbonate and Ca²⁺ transport in high-calcifying cells of E. huxleyi. Bicarbonate availability had significant and rapid effects on pH_i in exponential- but not in stationary-phase cells. 4′, 4′-Diisothiocyanostilbene-2,2′-disulfonic acid did not block bicarbonate uptake from the external medium by exponential-phase cells. Inorganic carbon utilization by exponential- and stationary-phase cells of Emiliania huxleyi was investigated using a pH drift technique in a closed system. Light-dependent alkalization of the medium by stationary-phase cells resulted in a final pH of 10.1 and was inhibited by dextran-bound sulphonamide, an inhibitor of external carbonic anhydrase. Exponential-phase cells did not generate a pH drift. Overall, the results suggest that for high-calcifying cultures of E. huxleyi the predominant pathway of inorganic carbon utilization differs in exponential and stationary phase cells of the same culture.     

Influence of galls of Phanacis taraxaci on carbon partitioning within common dandelion, Taraxacum officinale

We examined how leaf galls, induced by the cynipid wasp Phanacis taraxaci, influence the partitioning of photoassimilates within the host, the common dandelion, Taraxacum officinale. Galled and ungalled plants were exposed to ¹⁴CO₂ and the labelled photoassimilates accumulating within galls and other parts of the host were measured. During the growth phase of the gall they were physiological sinks for photoassimilates, accumulating 9% to 70% of total carbon produced by the host, depending upon the number of galls per plant. High levels of ¹⁴C assimilation in the leaves of galled plants compared to controls, suggest that galls actively redirect carbon resources from unattacked leaves of their host plant. This represents a significant drain on the carbon resources of the host, which increases with the number and size of galls per plant. Active assimilation of ¹⁴C by the gall is greatest in the growth phase and is several orders of magnitude lower in the maturation phase. This finding is consistent with physiological and anatomical changes that occur during the two phases of gall development and represents a key developmental strategy by cynipids to ensure adequate food resources before larval growth begins.     

Growth and carbon utilization by sprouted propagules of two species of submersed rooted aquatic plants grown in darkness

Hydrilla verticillata (L. f.) Royle tubers from monoecious plants andPotamogeton gramineus L. winter buds were sprouted and allowed to grow in the dark for 120 days. We measured plant length and counted the number of leaves at 2–3 day intervals.Hydrilla grew most rapidly during the first 16–17 days andPotamogeton grew most rapidly during the first 16–25 days. Measurement of propagule carbon content over time indicated that cessation of rapid growth coincided with depletion of tuber carbon by one-half forHydrilla. ForPotamogeton, growth was reduced after 16 to 25 days while the winter bud C half-life was 37 days. Calculations indicated thatHydrilla mobilized 49% andPotamogeton 39% of the initial propagule carbon to support growth. In a second experiment, in which plants were grown in substrate the plants grew taller and produced slightly more leaves per plant.Potamogeton removed from darkness after specified time periods, and allowed to grow for 21 days in a greenhouse recovered from 20–30 days in the dark. Similarly treatedHydrilla plants recovered from up to 80 days in the dark.Potamogeton had mobilized 79% of initial C by the time it was unable to recover from the dark treatment. Combined results for both species indicate that the majority of propagule C was utilized in the first 16 to 30 days following sprouting. In conjunction with an understanding propagule sprouting requirements, this information will be useful in the timing of application for management techniques. The U.S. Government right to retain a non-exclusive, royalty-free licence in and to any copyright is acknowledged. The U.S. Government right to retain a non-exclusive, royalty-free licence in and to any copyright is acknowledged.     

A long-term perspective of dissolved organic carbon transport in Sycamore Creek, Arizona, USA

Dissolved organic carbon (DOC) dynamics were examined over five years (1989–1993) in Sycamore Creek, a Sonoran Desert stream, specifically focusing on DOC concentration in surface and hyporheic waters, and rates of export. In 1989 and 1990, the years of lowest stream discharge (0.08 and 0.04 m³ s^–1 annual mean of daily discharge, respectively), DOC was high, averaging 7.37 and 6.22 mgC l^–1 (weighted annual means). In contrast, from 1991 through 1993, a period of increased flow (1.1, 1.2 and 4.3 m³ s^–1), concentration was significantly lower (P<0.001) with annual mean concentrations of 3.54, 3.49 and 3.39 mgC l^–1. Concentration exhibited little spatial variation between two sampling stations located 6 km apart along the mainstem or between surface and hyporheic waters. Annual export of DOC from Sycamore Creek varied 100-fold over the five-year period from a mean rate of only 24 kgC d^–1 in 1990 to 2100 kgC d^–1 in 1993. Ninety percent of DOC was exported by flows greater than 2.8 m³ s^–1, and 50% during flows greater than 27 m³ s^–1; flows of 2.8 and 24 m³ s^–1 occurred only 9 and 1% of the time. The export of organic matter in Sycamore Creek appears to be coupled to El Niño-Southern Oscillation phenomena. The years of highest export, 1991–1993, had El Niño conditions while 1989 and 1990 had medial conditions.     

Effects of the diatom-Emiliana huxleyi succession on photosynthesis, calcification and carbon metabolism by size-fractioned phytoplankton

Changes in the species composition, photosynthesis, calcification and size-fractionated carbon metabolism by natural phytoplankton assemblages were monitored in three mesocosms under different nutrient conditions during May 1993. In the 3 enclosures, the decline of the diatom-dominated assemblages was followed by the development of a bloom of the coccolithoporid Emiliania huxleyi. Highest growth of E. huxleyi was observed in the mesocosm with a high N : P ratio, suggesting this species is a good competitor at low phosphate concentrations. The transition from diatom- to E. huxleyi-dominated assemblages brought about a sharp reduction of the phytoplankton standing stock and carbon-specific photosynthetic rate. The relative contribution of the smaller size fraction to total photosynthesis increased as the succession progressed. Calcification rate and E. huxleyi cell-specified calcite production were highest during the early stages of development of the E. huxleyi bloom. Distinct changes in the patterns of ¹⁴C allocation into biomolecules were noticed during the diatom-E. huxleyi succession. The diatom-dominated assemblage showed high relative ¹⁴C incorporation into low molecular weight metabolites (LMWM), whereas proteins and, specially, lipids accounted for the largest proportion of carbon incorporation in the E. huxleyi bloom. The patterns of photoassimilated carbon metabolism proved to be strongly dependent on cellular size, as protein relative synthesis was significantly higher in the smaller than in the larger size fraction, irrespective of the nutrient regime and the successional stage. These results are discussed in relation to the ecological and physiological features of small phytoplankton.     

Ecophysiology and distribution of the endemic leafless spurgeEuphorbia aphylla and the introducedE. tirucalli (Euphorbiaceae,Euphorbia sect.Tirucalli) in the Canary Islands

The leafless spurgeEuphorbia aphylla (Euphorbiaceae), an endemic species restricted to several of the Canary Islands where it inhabits coastal and arid localities, expresses Crassulacean Acid Metabolism (CAM) when it is subjected to summer drought. A flexible CAM is consistent with the general ecology of the species. It is the only member of sect.Tirucalli native to the Canary Islands and is at the north-western edge of the section's biogeographical range. The other members of the section have a paleotropic distribution and are found throughout Africa. Many of them are regarded as obligate CAM plants, includingE. tirucalli which was used as a comparison in ecophysiological experiments examining the response ofE. aphylla to drought and temperature.     

Stable carbon isotopes as indicators of increased water use efficiency and productivity in white spruce (Picea glauca (Moench) Voss) seedlings

The relationship among water use efficiency (WUE), productivity and carbon isotopic composition (δ¹³C) in white spruce (Picea glauca (Moench) Voss) seedlings was investigated. Sixteen hundred seedlings representing 10 controlled crosses were planted in the field in individual buried sand-filled cylinders. The soil water content in the cylinders was measured using time domain reflectometry over two growing seasons and seedling water use determined by water balance. Two watering treatments were imposed: irrigation and dry land. There was significant (1.6–2.0%c) genetic variation in needle δ¹³C. Ranking of crosses in terms of δ¹³C was generally maintained over watering treatments and there was not a significant genetic versus environmental interaction. There was a positive correlation between δ¹³C and both intrinsic and long-term WUE (more positive δ¹³C with increased WUE) and between δ¹³C and productivity, suggesting a correlation due to variation in photosynthetic capacity. Root to shoot ratios did not increase in water-stressed plants, indicating that responses to drought were primarily at the level of gas exchange, rather than through morphological changes. Our results indicate that it should be possible to use δ¹³C as a surrogate for WUE and to select white spruce genotypes for high WUE without compromising yield.     

A meta-analysis of leaf gas exchange and nitrogen in trees grown under elevated carbon dioxide

The response of trees to rising atmospheric CO₂ concentration ([CO₂]) is of concern to forest ecologists and global carbon modellers and is the focus of an increasing body of research work. I review studies published up to May 1994, and several unpublished works, which reported at least one of the following: net CO₂ assimilation (A), stomatal conductance (g_s), leaf dark respiration (R_d) leaf nitrogen or specific leaf area (SLA) in woody plants grown at <400 μmol mol^?1 CO₂ or at 600–800 μmol mol^?1 CO₂. The resulting data from 41 species were categorized according to growth conditions (unstressed versus stressed), length of CO₂ exposure, pot size and exposure facility [growth chamber (GC), greenhouse (GH), or open-top chamber (OTC)] and interpreted using meta-analytic methods. Overall, A showed a large and significant increase at elevated [CO₂] but length of CO₂ exposure and the exposure facility were important modifiers of this response. Plants exposed for < 50 d had a significantly greater response, and those from GCs had a significantly lower response than plants from longer exposures or from OTC studies. Negative acclimation of A was significant and general among stressed plants, but in unstressed plants was influenced by length of CO₂ exposure, the exposure facility and/or pot size. Growth at elevated [CO₂] resulted in moderate reductions in gs in unstressed plants, but there was no significant effect of CO₂ on gs in stressed plants. Leaf dark respiration (mass or area basis) was reduced strongly by growth at high [CO₂] > while leaf N was reduced only when expressed on a mass basis. This review is the first meta-analysis of elevated CO₂ studies and provides statistical confirmation of several general responses of trees to elevated [CO₂]. It also highlights important areas of continued uncertainty in our understanding of these responses.     

Carbon isotope fractionation in tree ring early and late wood in relation to intra-growing season water balance

We determined the stable carbon isotope composition (δ^1.3C) of cellulose extracted from early and late wood in Douglas fir [Pseudotsuga menziexii (Mirb.) Franco] tree rings. Data were obtained for the period 1962 to 1981, at the start of which the trees were 20 years old. A water balance model was used to calculate daily stand transpiration and water deficit. The model incorporates site factors (soil water availability, slope and aspect) and environmental variables (solar radiation, air temperature and rainfall). There was far greater variability in late wood than in early wood δ^1.3C. In wet years, late wood δ^1.3C was significantly lighter (by as much as 2δ) than early wood δ^1.3C but in dry years this difference was reversed. Differences between spring and summer cumulative transpiration accounted for almost 60δ of the variability in differences between early and late wood δ^1.3C. We found excellent correspondence between summer cumulative transpiration and late wood δ^1.3C, with estimates of transpiration accounting for up to 93% of the variability in δ^1.3C. Correlations between early wood δ^1.3C and spring transpiration were generally poor (r2<0.4), but we were able to identify those exceptional years in which there had been a very dry spring. Our results indicate that, while tree ring δ^1.3C correlates reasonably well with basal area increment, it is a far better indicator of inter- and intra-annual variability in water availability than radial growth.