Photoadaptation of carboxylating enzymes and photosynthesis during a spring bloom

Properties of the light saturation curve of photosynthesis and ribulose-1,5-bisphosphate carboxylase (RuBPC) activity are shown to change qualitatively in a natural population of marine phytoplankton during a spring bloom. Evidence is presented to show that these changes constitute photoadapative responses to increasing irradiance. As irradiance increased during the bloom, both the level of light-saturated photosynthesis (P_m) and the initial slope of the light saturation curve (α = photosynthetic efficiency) increased whether those parameters were normalized to chlorophyll a concentration (P_m^b, α^b) or to cell numbers (P_m^c, α^c). The magnitudes of these changes were such that I_k (= P_m/α, the photoadaptation parameter) did not change, but I_m, the light intensity at which photosynthesis becomes saturated, increased. RuBPC activity, both chlorophyll a (RuBPC^b) and cell number normalized (RuBPC^c), also increased during the bloom. We suggest that these adaptations were achieved by simultaneously increasing the number of photosynthetic units, proportionately decreasing the photosynthetic unit size, and increasing both the concentrations of the enzymes of the dark reactions and possibly also of photosynthetic electron transport components.We also observed diminished levels of photoinhibition in the high light adapted cells late in the bloom and have suggested that this was a consequence of the same suite of physiological changes.In situ carbon fixation per cell increased during the bloom whereas no change occurred in this parameter when normalized to chlorophyll a concentration. Although these photoadaptive responses thus permitted carbon to be fixed in situ more rapidly per cell, at a constant efficiency with respect to investment of energy in the photosynthetic apparatus, they did not result in a change in growth rate. Based on consideratios of the role of time scale in physiological adaptation, however, it is suggested that the observed alterations in photosynthesis with increasing irradiance might permit a cell to more rapidly fill an energy quota for division, possibly an advantage in a mixing environment in which energy is patchily distributed, both spatially and temporalyy.Phosphoenolpyruvate carboxylase activity when normalized to chlorophyll a (PEPC^b) did not change during the bloom while chlorophyll a normalized dark carbon fixation decreased sharply and was quantitatively small compared to PEPC^b. On this basis and considering that RuBPC^b increased during the bloom, it is suggested that, although PEPC may be involved in dark carbon fixation, its most important quantitative role is probably an indirect one in light dependent photosynthesis.We have also considered the relevance of laboratory results on photoadaptation to interpretations of field studies and have suggested that batch culture studies must be treated with caution but that turbidistat and semi-continuous methods provide reasonable simulations of natural conditions.     

Temporal variability in phytodetritus and megabenthic activity at the seabed in the deep Northeast Atlantic

We report a ten-year study of the abundance and activity of megabenthos on the Porcupine Abyssal Plain, northeast Atlantic, together with observations on the occurrence of phytodetritus at the deep-sea floor (4850 m). Using the Southampton Oceanography Centre time-lapse camera system, ‘Bathysnap’, we have recorded a radical change in the abundance and activity of megabenthos between the two periods of study (1991–1994 and 1997–2000). In 1991–1994, the larger megabenthos occurred at an abundance of c. 71.6/ha and were dominated by large holothurians. In addition, there were very substantial populations of smaller megabenthic ophiuroids (c. 4979/ha). Together, the total megabenthos are estimated to track over some 17 cm²/m²/d (exploiting 100% of the surface of the seabed in c. 2.5 years). In 1997–2000, the larger megabenthos increased to an abundance of c. 204/ha and were joined by exceptional numbers of a small holothurian species (Amperima rosea, 6457/ha) and ophiuroids (principally Ophiocten hastatum, 53,539/ha). The total megabenthos population was tracking at an estimnated rate of c. 247 cm²/m²/d (exploiting 100% of seabed in just 6 weeks). Coincident with these increases in the abundance and activity of the megabenthos, there were apparently no mass depositions of aggregated phytodetritus to the seabed in the summers of 1997–1999. Mass occurrences of phytodetritus had been noted during the summer months of the three years previously studied (1991, 1993 and 1994), with covering between 50 and 96% of the sediment surface. There is a statistically significant (p<0.02) negative correlation between maximum extent of this seabed cover of phytodetritus and seabed tracking by megabenthos. Additional studies [Lampitt et al., Progr. Ocean. 50 (2001)], indicate that there were no substantial changes in surface ocean primary productivity, in export flux, or in the composition of the flux that might otherwise account for the apparent absence of observable concentrations of phytodetritus during the summers of 1997–1999. We postulate that the marked increase in megabenthic tracking activity resulted in the removal (via consumption, disaggregation, burial etc.) of the bulk of the incoming phytodetrital flux during these years. A simple conceptual model, based on the apparent phytodetrital fluxes observed in 1991 and 1993, suggests that the megabenthos tracking rates estimated for 1997–1999 are sufficient to account for near-total removal of this flux. However, we are not able to estimate other processes removing phytodetritus (i.e. other elements of the benthos) that may also have increased between 1991–1994 and 1997–1999. Other independent studies [e.g. Ginger et al., Progr. Ocean. 50 (2001)] of flux constituents support the possibility that just a few species of megabenthos (e.g. A. rosea, and O. hastatum) could well have consumed a major proportion of the incoming flux and so substantially modified the composition of the organic matter available to other components of the benthos.     

The impact of mesoscale eddies on plankton dynamics in the upper ocean

A coupled QuasiGeostrophic mixed-layer ECOsystem model (QGECO) is used to investigate the impact of the underlying mesoscale eddy field on the spatial and temporal scales of biological production and on overall rates of primary productivity. The model exhibits temporal trends in the biological and physical fields similar to those observed in the North Atlantic; i.e. the mixed layer shallows in spring causing a rapid increase in phytoplankton concentrations and a corresponding decline in nutrient levels. Heterogeneity is produced in the mixed layer through Ekman pumping velocities resulting from the interaction of windstress and surface currents. This variability impacts on biological production in two ways. Firstly, spatial variations in the depth of the mixed layer affect the photosynthetically active radiation (PAR) availability and hence production rates, and secondly, eddy enhanced exchange between the surface water and those at depth bring additional nutrients into the euphotic zone. These processes result in significant spatial and temporal heterogeneity in the ecosystem distributions.Investigation of the spatial heterogeneity of the biological system finds variability to be significantly greater than that of the mixed layer. The relationship between the eddy field and the ecosystem is investigated. The structure and correlation of the biogeochernical fields change with time. The biological fields are found to have a shorter horizontal scale, but whiter spectrum than the underlying eddy field.Overwinter conditions are found to have a profound effect on the variability, size and timing of the following spring bloom event. Variations in the nitrate levels are primarily responsible for the variability in the biological system in the first year. In subsequent years the variation in the overwintering population is found to be dominant.     

Errors in recent ocean tide models: possible origin and cause

Recently, the TOPEX/POSEIDON Science Working Team has recommended the FES95.2.1 and CSR3.0 ocean tide models for reprocessing the TOPEX/POSEIDON Geophysical Data Records. Without doubt, the performance of these models, especially in the deep oceans, is excellent. However, from a comparison of these hydrodynamically consistent models with the purely empirical DW3.2 and DEOS96.1 models, it appears that FES95.2.1 and CSR3.0 are affected by basin boundary related errors which are caused by the basin-wise solution procedure of the FES ocean tide model series. In their turn, the empirical DW3.2 and DEOS96.1 models seem to suffer from significant errors in the Antarctic seas due to the seasonal growth and decay of Antarctic sea ice. Also, bathymetry-induced differences were found between the hydrodynamically consistent models and the empirical models. Concerning these differences, TOPEX/POSEIDON and ERS-1 crossover statistics unfortunately do not provide conclusive results on which models are in error.     

Cross-Calibration and Long-Term Monitoring of the Microwave Radiometers of ERS, TOPEX, GFO, Jason, and Envisat

The radiometers on board the satellites ERS-1, TOPEX/Poseidon, ERS-2, GFO, Jason-1, and Envisat measure brightness temperatures at two or three different frequencies to determine the total columnal water vapor content and wet tropospheric path delay, a major correction to the altimeter range measurements. In order to asses the long-term stability of the path delay, the radiometers are calibrated against vicarious cold and hot references, against each other, and against several atmospheric models. Four of these radiometers exhibit significant drifts in at least one of the channels, resulting in yet unmodeled errors in path delay of up to 1 mm/year, thus limiting the accuracy at which global sea level rise can be inferred from the altimeter range measurements.     

A Sensitivity Analysis of the Waterline Method of Constructing a Digital Elevation Model for Intertidal Areas in ERS SAR scene of Eastern England

A sensitivity analysis of the waterline method of constructing a Digital Elevation Model (DEM) of an intertidal zone using remote sensing and hydrodynamic modelling is described. Variation in vertical height accuracy as a function of beach slope is investigated using a set of nine ERS Synthetic Aperture Radar (SAR) images of the Humber/Wash area on the English east coast acquired between 1992 and 1994. Waterlines from these images are heighted using a hydrodynamic tide-surge model and interpolated using block kriging. On 1:500 slope beaches, an average block height estimation standard deviation of 18–22 cm is achieved. This rises to 27 cm on 1:100 slope beaches, and 32 cm on 1:30 slope beaches. The average heighting error at different slopes is decomposed into components due to waterline heighting error, inadequate sensor resolution and interpolation inaccuracy. It is shown that, at 1:500 slope, waterline heighting error and interpolation inaccuracy are the main error sources, whilst at 1:30 slope, errors due to inadequate sensor resolution become dominant. The ability of the technique to generate intertidal DEMs for almost the entire coastal zone in a complete ERS SAR scene covering 100×100 km is demonstrated.