Description: Marine life is controlled by multiple physical and chemical drivers and by diverse ecological processes. Many of these oceanic properties are being altered by climate change and other anthropogenic pressures. Hence, identifying the influences of multifaceted ocean change, from local to global scales, is a complex task. To guide policy-making and make projections of the future of the marine biosphere, it is essential to understand biological responses at physiological, evolutionary and ecological levels. Here, we contrast and compare different approaches to multiple driver experiments that aim to elucidate biological responses to a complex matrix of ocean global change. We present the benefits and the challenges of each approach with a focus on marine research, and guidelines to navigate through these different categories to help identify strategies that might best address research questions in fundamental physiology, experimental evolutionary biology and community ecology. Our review reveals that the field of multiple driver research is being pulled in complementary directions: the need for reductionist approaches to obtain process-oriented, mechanistic understanding and a requirement to quantify responses to projected future scenarios of ocean change. We conclude the review with recommendations on how best to align different experimental approaches to contribute fundamental information needed for science-based policy formulation.

Description: As part of the PeECE II mesocosm project, we investigated the effects of pCO2 levels on the initial step of heterotrophic carbon cycling in the surface ocean. The activities of microbial extracellular enzymes hydrolyzing 4 polysaccharides were measured during the development of a natural phytoplankton bloom under pCO2 conditions representing glacial (190 µatm) and future (750 µatm) atmospheric pCO2. We observed that (1) chondroitin hydrolysis was variable throughout the pre-, early- and late-bloom phases, (2) fucoidanase activity was measurable only in the glacial mesocosm as the bloom developed, (3) laminarinase activity was low and constant, and (4) xylanase activity declined as the bloom progressed. Concurrent measurements of microbial community composition, using denaturing-gradient gel electrophoresis (DGGE), showed that the 2 mesocosms diverged temporally, and from one another, especially in the late-bloom phase. Enzyme activities correlated with bloom phase and pCO2, suggesting functional as well as compositional changes in microbial communities in the different pCO2 environments. These changes, however, may be a response to temporal changes in the development of phytoplankton communities that differed with the pCO2 environment. We hypothesize that the phytoplankton communities produced dissolved organic carbon (DOC) differing in composition, a hypothesis supported by changing amino acid composition of the DOC, and that enzyme activities responded to changes in substrates. Enzyme activities observed under different pCO2 conditions likely reflect both genetic and population-level responses to changes occurring among multiple components of the microbial loop.

Description: The carbon and nitrogen content of transparent exopolymer particles (TEP) was determined and related to the concentration of TEP as quantified by a colorimetrical method. TEP were produced in the laboratory from dissolved precursors by laminar or turbulent shear. Dissolved precursors were obtained by 0.2 µm filtration from diatom cultures, with or without nutrient reduction, and from natural diatom populations. The relationship between carbon and TEP was significant, linear and species-specific. Carbon concentration of TEP derived from this relationship concurred with previous findings. Shortage of silicic acid or nitrate in the culture media had no effect on the carbon content of TEP. Molar C:N ratios of TEP were above the Redfield ratio, with a mean value of 26. It is suggested that the nitrogen fraction of TEP can be explained by adsorption of dissolved organic nitrogen (DON) onto TEP. Based on the newly established relationship, concentrations of TEP-derived carbon (TEP-C) were calculated for the Baltic Sea, the coastal Pacific, the North East Atlantic and the Northern Adriatic Sea.

Description: The role of TEP (Transparent Exopolymer Particles) in the flocculation of a diatom bloom was studied under controlled conditions in a mesocosm. The concentration of TEP increased exponentially during growth, flocculation and senescence of the bloom. Aggregation began dominating the particle dynamics of TEP during the early growth phase of the bloom, several days prior to the appearance of large flocs and nutrient depletion. TEP aggregated with themselves and with phytoplankton due to the high stickiness of TEP, but phytoplankton was not observed to aggregrate with itself. The production of TEP, estimated from changes in concentration, did not increase after nutrients were depleted. The concentration of TEP was a linear function of chl a and particulate organic carbon (POC), indicating that production of TEP was linked to growth rather than standing stocks of phytoplankton. The ratio between TEP and phytoplankton appeared to be one of the factors determining the onset of the flocculation of the bloom. The concentration of TEP may have been decreased by bacterial degradation. Bacterial degradation of TEP may explain the low TEP to chl a values, the decrease in stickiness of particles as the bloom progressed, and the retarded onset of flocculation.

Description: Flows of the major biogeochemical elements (C, N, P, Si) and of transparent exopolymer particles (TEP) were traced during a bloom of a natural assemblage of marine diatoms in a mesocosm (l m(3)) to determine whether the exudation and subsequent gelation of carbon-rich phytoplankton exopolymers can account for the formation and potential export of carbon in excess of that predicted by Redfield ratios. Exponential growth of the phytoplankton community in the mesocosm extended for 10 d until nitrate concentration fell below detection and concentrations of dissolved inorganic and particulate organic nitrogen and phosphorus remained stable. Tight covariation of particulate organic elements occurred as long as nutrients were replete. But, after nitrate depletion, decoupling of carbon dynamics from that of nitrogen and phosphorus was observed, with a large flow of carbon into TEP An uptake of 72% more dissolved inorganic carbon (DIC) than inferred from nitrate supply and Redfield stoichiometry (referred to as carbon overconsumption) occurred during the study, largely during the postbloom phase, and was almost entirely traced to the particulate organic matter (POM) pool. Marine snow (aggregates 〉0.5 mm) appeared at the onset of nitrate depletion and coincided with rapid increase in TEP concentrations. Elemental composition of marine snow differed from the Redfield ratio by an enrichment in carbon and a depletion in phosphorus relative to nitrogen. It is suggested that sinking of TEP-rich marine snow could be a possible mechanism for export of carbon above calculations that are based on the Redfield stoichiometry.

Description: Observations that the majority of silica dissolution occurs within the upper 200 m of the ocean, and that sedimentation rates of diatom frustules generally do not decrease significantly with depth, suggested reduced dissolution rates of diatoms embedded within sinking aggregates. To investigate this hypothesis, silica dissolution rates of aggregated diatom cells were compared to those of dispersed cells during conditions mimicking sedimentation below the euphotic zone. Changes in the concentrations of biogenic silica, silicic acid, cell numbers, chlorophyll a and transparent exopolymer particles (TEP) were monitored within aggregates and in the surrounding seawater (SSW) during two 42-day experiments. Whereas the concentration of dispersed diatoms decreased over the course of the experiment, the amount of aggregated cells remained roughly constant after an initial increase. Initially only 6% of cells were aggregated and at the end of the experiment more than 60% of cells were enclosed within aggregates. These data imply lower dissolution rates for aggregated cells. However, fluxes of silica between the different pools could not be constrained reliably enough to unequivocally prove reduced dissolution for aggregated cells.

Description: Zwischen 7. September 1983 und 19. Januar 1984 wurden in ca. wöchentlichem Abstand Proben an einer Station in der westlichen Kieler Bucht genommen, um die Entwicklung und Sedimentation der Herbstblüten zu untersuchen. In dieser Arbeit wurde die Phytoplanktonsukzession in Abhängigkeit von der hydrographischen Situation, der Nährsalzkonzentration und der Lichtbedingungen dargestellt. Im Herbst 1983 fanden in der Kieler Bucht zwei ausgeprägte Blüten statt. Die erste Blüte stellten C.tripos und C.fusus; sie erreichte ihr Maximum von 8,6gPPC/m² am 5.10. Die zweite Blüte wurde von den Diatomeen Thalassiosira decipiens und Chaetoceros curvisetus dominiert; ihr Maximum lag mit 6.5gPPC/m² am 28.11.83. Für den Aufbau der Ceratienblüte herrschten nach Nährsalzeintrag in die vom Somer nährsalzverarmte Oberflächenschicht optimale Entstehungsbedingungen. Hohe Primärproduktion (0,8-1gC/m²/d) und hohe Assimilationszahlen (über 6) während der Blüte bewiesen das gute Wachstum der Ceratien. Als Ursache der Sukzession vom Sommerplankton zu Ceratien werden der geringe Freßdruck auf die Mobilität der Ceratien diskutiert. Niedrige Turbulenzen werden als entscheidendes Kriterium für den Erfolg von C.tripos gegenüber C.fusus im Herbst 1983 angeführt. Gründe für die Sedimentation der Ceratienblüte sind aus direkten Beziehungen zwischen Umweltbedingungen und Wachstum nicht ersichtlich da Nährsalze und Lichtangebot ausreichend waren, und weder interspezifische Konkurrenz noch Freßfeinde vorhanden waren. Es wird vorgeschlagen die Gründe für das plötzliche Verschwinden der Ceratienblüte in der Organismen zu suchen. Intrinsischer Wechsel von vegetativer und geschlechtlicher Vermehrung zusammen mit unterschiedlich starker Umweltabhängigkeit dieser Vermehrungsmodi können viele Phänomene, die im Zusammenhang mit Ceratien berichtet worden sind, erklären. Der Aufbau der Diatomeen nah der fast vollständigen Sedimentation der Ceratienblüte wurde trotz niedriger Einstrahlungsintensität durch das Entstehen einer erneuten Schichtung ermöglicht. Trotz sehr hoher Lichtausnutzung der Diatomeen, war die Primärproduktion aufgrund niedriger Einstrahlungstrahlung gering (0,5g/m²/d). Für die Sedimentation der Diatomeen war die rapide Verschlechterung des Lichtklimas Hauptursache. Die Stickstoffkonzentrationen lagen in einem Bereich, in dem sie als Streßfaktor angesehen und für die Sedimentation mitverantwortlich gemacht werden können.

Description: Optical particle measurements are emerging as an important technique for understanding the ocean carbon cycle, including contributions to estimates of their downward flux, which sequesters carbon dioxide (CO2) in the deep sea. Optical instruments can be used from ships or installed on autonomous platforms, delivering much greater spatial and temporal coverage of particles in the mesopelagic zone of the ocean than traditional techniques, such as sediment traps. Technologies to image particles have advanced greatly over the last two decades, but the quantitative translation of these immense datasets into biogeochemical properties remains a challenge. In particular, advances are needed to enable the optimal translation of imaged objects into carbon content and sinking velocities. In addition, different devices often measure different optical properties, leading to difficulties in comparing results. Here we provide a practical overview of the challenges and potential of using these instruments, as a step toward improvement and expansion of their applications.