Characterization of disjunct populations of Zostera marina (eelgrass) from California: genetic differences resolved by restriction-fragment length polymorphisms

Comparative restriction-fragment analysis was used to analyze the nuclear ribosomal DNA, and alcohol dehydrogenase-1 loci of Zostera marina L., for variation within and among populations. Eelgrass is a perennial marine flowering plant that is widespread and ecologically significant throughout the temperate northern hemisphere. A chemical method was developed to obtain restriction-quality DNA without CsCl fractionation from experimentally relevant quantities of seagrass tissues (0.5 to 1.0 g). The yield was 25 g g^-1 fresh weight. The three morphologically distinct forms of Z. marina from disjunct populations examined in this study were found to be genetically distinct; morphologically similar populations were indistinguishable genetically. Genetic distinction also correlated with habitat depth, as subtidal and intertidal populations were clearly divergent. Homologous probes for the 17S and 28S ribosomal DNA genes were used to map 24 restriction sites on the rDNA repeat of Z. marina, which was determined to be about 14 kb in length. At least 1 length mutation and 5 restriction-site changes were identified that distinguished Z. marina populations from San Diego and Monterey Bay (Del Monte Beach) from Z. marina populations from Elkhorn Slough and Tomales Bay. Estimated sequence variation (100×p) between eelgrass populations ranged from 0.00 to 0.69. Individual plants were observed to contain as many as four different rDNA-repeat length variants. The mean number of rDNA-repeat length variants per individual in Z. marina was about two. Intrapopulation variation in rDNA-repeat type was observed in only one individual from the Tomales Bay population.     

Sediment ammonium availability and eelgrass (Zostera marina) growth

The interaction of sediment ammonium (NH ₄ ⁺ ) availability and eelgrass (Zostera marina L.) growth, biomass and photosynthesis was investigated using controlled environment and in-situ manipulations of pore water ammonium concentrations. Sediment diffusers were used to create pore water diffusion gradients to fertilize and deplete ammonium levels in sediments with intact eelgrass rhizospheres. Between October, 1982 and September, 1983 controlled environment experiments using plants from shallow (1.3 m) and deep (5.5 m) stations in a Great Harbor, Woods Hole, Massachusetts, USA eelgrass meadow along with in-situ experiments at these stations provided a range of sediment ammonium concentrations between 0.1 and 10 mM (adsorbed+interstitial NH ₄ ⁺ ). The results of the in-situ experiments indicate that nitrogen limitation of eelgrass growth does not occur in the Great Harbor eelgrass meadow. A comparison of NH ₄ ⁺ regeneration rates and eelgrass nitrogen requirements indicates an excess of nitrogen supply over demand and provides an explanation for the lack of response to the manipulations. Results of controlled environment experiments combined with in-situ results suggest that sediment ammonium pool concentrations above approximately 100 mol NH ₄ ⁺ per liter of sediment (interstitial only) saturate the growth response of Zostera marina.     

Effects of anaerobiosis on root metabolism of Zostera marina (eelgrass): implications for survival in reducing sediments

The temperate seagrass Zostera marina L. typically grows in highly reducing sediments. Photosynthesis-mediated O₂ supplied to below-ground tissues sustains aerobic respiration during photosynthetic periods. Roots, however, experience daily periods of anoxia and/or hypoxia at night and under conditions that reduce photosynthesis. Rhizosphere cores of Z. marina were collected in August 1984 from Great Harbor, Massachusetts, USA. We examined short-term anaerobic metabolism of [U-¹⁴C]sucrose in excised roots and roots of intact plants. Under anaerobic conditions roots showed appreciable labeling of CO₂, ethanol and lactate, and slight labeling of alanine and other metabolites. Over 95% of the ¹⁴C-ethanol was recovered in the root exudate. Release of other metabolites from the roots was minimal. Ethanol was also released from hypoxic/anoxic roots of intact plants and none of this ethanol was transported to the shoot under any experimental conditions. Loss of ethanol from roots prevented tissue levels of this phytotoxin from increasing during anaerobiosis despite increased synthesis of ethanol. Anaerobic metabolism of [U-¹⁴C]glutamate in excised roots led to appreciable labelling of -aminobutyrate, which was known to accumulate in eelgrass roots. Roots recovered to fully aerobic metabolism within 4 h after re-establishment of aerobic conditions. The contributions of these root metabolic responses to the ability of Z. marina to grow in reducing marine sediments are related to light-regulated interactions of shoots and roots that likely dictate depth penetration, distribution and ecological success of eelgrass.     

Photosynthetic characteristics of Prochloron sp./ascidian symbioses

The prokaryotic green alga Prochloron sp. (Prochlorophyta) is found in symbiotic association with colonial didemnid ascidians that inhabit warm tropical waters in a broad range of light environments. We sought to determine the light-adaptation features of this alga in relation to the natural light environments in which the symbioses are found, and to characterize the temperature sensitivity of photosynthesis and respiration of Prochloron sp. in order to assess its physiological role in the productivity and distribution of the symbiosis. Colonies of the host ascidian Lissoclinum patella were collected from exposed and shaded habitats in a shallow lagoon in Palau, West Caroline Islands, during February and March, 1983. Some colonies from the two light habitats were maintained under conditions of high light (2 200 E m^–2 s^–1) and low light (400 E m^–2 s^–1) in running seawater tanks. The environments were characterized in terms of daily light quantum fluxes, daily periods of light-saturated photosynthesis (H_sat), and photon flux density levels. Prochloron sp. cells were isolated from the hosts and examined for their photosynthesis vs irradiance relationships, respiration, pigment content and photosynthetic unit features. In addition, daily P:R ratios, photosynthetic quotients, carbon balances and photosynthetic carbon release were also characterized. It was found that Prochloron sp. cells from low-light colonies possessed lower chlorophyll a/b ratios, larger photosynthetic units sizes based on both reaction I and reaction II, similar numbers of reaction center I and reaction center II per cell, lower respiration levels, and lower P_max values than cells from high-light colonies. Cells isolated from low-light colonies showed photoinhibition of P_max at photon flux densities above 800 E m^–2 s^–1. However, because the host tissue attenuates about 60 to 80% of the incident irradiance, it is unlikely that these cells are normally photoinhibited in hospite. Collectively, the light-adaptation features of Prochloron sp. were more similar to those of eukaryotic algae and vascular plant chloroplasts than to those of cyanobacteria, and the responses were more sensitive to the daily flux of photosynthetic quantum than to photon flux density per se. Calculation of daily minimum carbon balances indicated that, though high-light cells had daily P:R ratios of 1.0 compared to 4.6 for low-light cells, the cells from the two different light environments showed nearly identical daily carbon gains. Cells isolated from high-light colonies released between 15 and 20% of their photosynthetically-fixed carbon, levels sufficient to be important in the nutrition of the host. Q₁₀ responses of photosynthesis and respiration in Prochloron sp. cells exposed briefly (15–45 min) to temperatures between 15° and 45°C revealed a discontinuity in the photosynthetic response at the ambient growth temperatures. The photosynthetic rates were found to be more than twice as sensitive to temperatures below ambient (Q₁₀=3.47) than to temperatures above ambient (Q₁₀=1.47). The Q₁₀ for respiration was constant (Q₁₀=1.66) over the temperature range examined. It appears that the photosynthetic temperature sensitivity of Prochloron sp. may restrict its distribution to warmer tropical waters. The ecological implications of these findings are discussed in relation to published data on other symbiotic systems and free-living algae.     

Light regulation of nitrate reductase in Ulya fenestrata (Chlorophyceae)

Observations were made on the behavior of nitrate reductase activity in the green alga Ulva fenestrata under controlled light:dark regimes. The activity of nitrate reductase (NR) was examined in response to normal seasonal photoperiods as well as in response to shortened or extended periods of darkness. NR activity exhibits a light-dependent diurnal rhythm under both normal summer and winter photoperiods, with a maximum in the early morning (2 to 2.5 h after the start of illumination). This peak of activity is followed by a lower steady-state level of activity which is sustained throughout the light period. There is a sustained minimal level of activity in darkness. The morning peak in activity is always observed as long as tissue is illuminated, irrespective of the previous light or dark treatments. As such, it appears that nitrate reductase activity in U. fenestrata is under circadian control. There is no major difference in the NR activity pattern between summer and winter plants, except that the peak activity values in winter plants are consistently much higher (5 times) than in summer plants. The study also suggests that illumination prior to the normal start of photoperiod triggers a different set of regulatory mechanisms, indicating that the physiological state of plants is important in dictating the NR activity response to illumination.Please address all correpondence and requests for reprints to G. J. Smith at his present address (Hopkins Marine Station)     

Scenarios of global bioenergy production: The trade-offs between agricultural expansion, intensification and trade

Increased future demands for food, fibre and fuels from biomass can only be met if the available land and water resources on a global scale are used and managed as efficiently as possible. The main routes for making the global agricultural system more productive are through intensification and technological change on currently used agricultural land, land expansion into currently non-agricultural areas, and international trade in agricultural commodities and processed goods. In order to analyse the trade-offs and synergies between these options, we present a global bio-economic modelling approach with a special focus on spatially explicit land and water constraints as well as technological change in agricultural production. For a global bioenergy demand scenario reaching 100 ExaJoule (EJ) until 2055 we derive a required rate of productivity increase on agricultural land between 1.2 and 1.4 percent per year under different land allocation options. A very high pressure for yield increase occurs in Sub-Saharan Africa and the Middle East, even without additional bioenergy demand. Moreover, we analyse the implicit values (shadow prices) of limited water resources. The shadow prices for bioenergy are provided as a metric for assessing the trade-offs between different land allocation options and as a link between the agricultural and energy sector.     

Ecotypic differentiation in the marine diatom Skeletonema costatum: influence of light intensity on the photosynthetic apparatus

Three genetically distinct clones of Skeletonema costatum (Grev.) Cleve were grown at 20°C under high (274 E m^-2 s^-1) and low (27 E m^-2 s^-1) light conditions and their photoadaptive photosynthetic responses compared. When all three clones were grown under low light, pigment analyses and fluorescence excitation spectra demonstrated that the accessory pigments, chlorophyll c and fucoxanthin, became more important in light-harvesting compared to chlorophyll a. Photosynthetic unit sizes increased for Photosystems I and II in low light, but photosynthesis vs irradiance characteristics were not reliable predictors of photosynthetic unit features. Fluorescence excitation spectra and photosynthesis vs irradiance (P-I) relationships indicated that changes in energy transfer occurred independent of changes in pigment content. Large increases in accessory pigment content were not accompanied by large increases in excitation from these pigments. Changes in energy transfer properties were as important as changes in PSU size in governing the photoadaptive responses of S. costatum. When the three clones were grown under identical conditions, each had a separate and distinct pattern of photoadaptation. Significant differences among clones were found for pigment ratios, photosynthetic unit sizes for Photosystems I and II and efficiency of energy transfer between pigments. These strikingly different photoadaptive strategies among clones may partially account for the great ecological success of the diatom species. This is the first quantitative investigation of the importance of both chlorophyll c and fucoxanthin to the adaptive responses of diatoms to light intensity, and represents the most complete characterization of the photoadaptive responses of a single species of marine phytoplankter to differences in light environment.     

Harvesting the sun: New estimations of the maximum population of planet Earth

The maximum population, also called Earth's carrying capacity, is the maximum number of people that can live on the food and other resources available on planet Earth. Previous investigations estimated the maximum carrying capacity as large as about 1 trillion people under the assumption that photosynthesis is the limiting process. Here we use a present state-of-the-art dynamic global vegetation model with managed planetary land surface, Lund-Potsdam-Jena managed Land (LPJmL), to calculate the yields of the most productive crops on a global 0.5° × 0.5° grid. Using the 2005 crop distribution the model predicts total harvested calories that are sufficient for the nutrition of 11.4 billion people. We define scenarios where humankind uses the whole land area for agriculture, saves the rain forests and the boreal evergreen forests or cultivates only pasture to feed animals. Every scenario is run in an extreme version with no allowance for urban and recreational needs and in two soft versions with a certain area per person for non-agricultural use. We find that there are natural limits of the maximum carrying capacity which are independent of any increase in agricultural productivity, if non-agricultural land use is accounted for. Using all land planet Earth can sustain 282 billion people. The save-forests-scenario yields 150 billion people. The scenario that cultivates only pasture to feed animals yields 96 billion people. Nevertheless, we should always have in mind that all our calculated numbers for the carrying capacity refer to extreme scenarios where humankind may only vegetate on this planet. Our numbers are considerably higher than the general median estimate of upper bounds of human population found in the literature in the order of 10 billion.     

Characterization of in vivo absorption features of chlorophyte,phaeophyte and rhodophyte algal species

Despite the plentiful diversity of macroalgae in coastal environments, few studies have examined the in vivo absorption features of common marine macrophytes. Here we report on results of a survey of 12 central California common intertidal and subtidal taxa, representing Chlorophyta, Phaeophyta and Rhodophyta carried out in the summer of 1988. Computer-assisted analyses were used to obtain fourth-derivative spectra from in vivo absorption spectra determined at room temperature to obtain spectral diagnostics for the different algal divisions and to provide a means to determine whether spectral features could be used to identify stress responses among these plants. Among the Chlorophyta, characteristic maxima for chlorophylls a and b were resolved in all species examined, and a spectral component attributable to siphonaxanthin-like carotenoid(s) was observed in two Ulva species and the coenocytic alga Derbesia marina. Representatives of the Phaeophyta were characterized by similar maxima for chlorophyll a and c, and for fucoxanthin. Among the Rhodophyta, maxima for chlorophyll a, as well as B- and R-type phycoerythrin were resolved. Differences in in vivo absorption features were detected for two tidal populations of Porphyra perforata and Mastocarpus papillatus. High-tidal thalli absorb less green light in regions characterized by phycoerythrin, but have enhanced carotenoid absorption compared with lowtidal thalli. Resolution of spectra by fourth-derivative analysis revealed significant differences in phycoerythrin and carotenoid contents. The spectral changes observed appear to reflect environmental and possibly populational characteristics of these algae. The spectral analyses described here provide robust, non-invasive means to characterize subtle responses of macroalgae to environment in ways not possible previously. Additional merits of these fourth-derivative analyses for use in environmental studies are discussed.This contribution is dedicated to the memory of Professor L. R. Blinks, a pioneer in photosynthesis research and algal physiology     

Climatic risks and impacts in South Asia: extremes of water scarcity and excess

Regional Environmental Change - This paper reviews the current knowledge of climatic risks and impacts in South Asia associated with anthropogenic warming levels of 1.5–4 °C...