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Tabelle A13: Explorative Faktorenanalyse des HAWIK-IV (Erstvorgabe HAWIK-III). The meadian
nitrate concentration for these large Beggiatoa was thus ca 130 ?M. Indizes des HAWIK-IV 6.3.9-
6.3.10 Indizes des HAWIK-III vs. The flux budget further indicates a netoutgassing of CO2 to the
atmosphere of about 50 g C m?2 a?1 in the SRB. Neben dem WLD-Index zeigen auch einige
Untertests im Mittel hohere Werte im HAWIK-IV. They continued their gliding motility even down
to -5?C and immediately recovered, without motility loss, from a transient freezing at that
temperature (Dunker et al., in press). Since the upper part of the lagoonal sediment in Ymerbukta
freezes solid in winter, it is an important observation that both Beggiatoa and sulfate reducing
bacteria from this environment survive freezing without or with only little detrimental effect on their
metabolic rates (R. The grazing rate and aggregation rate are noted as gr and agg, respectively. With
these improvements, the model will be suitable to investigate the ocean’s response to vari-ations in
the physical forcing in climate change model scenarios and to simulate biogeochemicalfluxes on a
larger scale than a few decades. 112 Page 123. The physiological response to iron limitation is
independent of the assumed mechanism for siliconuptake. The narrow 5 ?m Beggiatoa had highest
density at 1-2 cm depth while the wider 12 ?m Beggiatoa had maximum at 2-3 cm depth (Fig. 5A).
Due to their much larger individual filament size, the biomass of the 12 ?m filaments was overall
highest (Fig. 5B). The mean areal biomass was 0.167 g m-2 of which the 5 ?m wide filaments
accounted for 40%. 85 Page 87. In this study, theshallow tidal basin between the german island of
Sylt and the danish island of Romo, the Sylt-Romobight or List tidal basin, has been chosen as study
area because a huge amount of data and informationis available for this system (Gatje and Reise,
1998). Excretion is parameterized as a linear function of phytoplankton biomass with constant excre-
tion or loss rates, ?, for carbon, nitrogen, and silicon. In REcoM phytoplank-ton carbon, nitrogen,
and chlorophyll are decoupled (variable stoichiometry). The two narrowest size classes, 2-3 ?m and
4-5 ?m, were the most abundant and comprised 77% of all Beggiatoa filaments counted. Beggiatoa
occurred throughout the 4 cm deep oxidized zone (Fig. 5) and two distinct size classes prevailed.
Oxygen in the benthic layer is removed by oxicremineralization. Sensing of stimuli can be achieved
either by spatial sensing or by temporal sensing. We will not try to reproduce the observed transition
but test thelinear nonsaturable uptake as an alternative to Michaelis-Menten kinetics. Bern: Huber.
Tewes, U. (1991). Hamburg-Wechsler-Intelligenztest fur Erwachsene - Revision (HAWIE-R). After
the filaments stopped, the temperature was lowered 54 Page 56. At temperatures above 5-10?C the
filaments glide at a velocity of 2-4 ?m s-1 and effectively migrate within their preferred 73 Page 75.
Degradation of detritus carbon and nitro-gen is implemented as a temperature dependent linear
function of detritus concentrations assumingthat this process only depends on substrate availability
and that the abundance of the decomposingorganisms linearly increases with detritus concentrations
with a constant degradation rate. This concentration gradient leads to a steady import of detritusand
extracellular organic matter into the bight. Intelligence, 33, 613-621. Kaufman, A. S. (1992).
Evaluation of the WISC-III and WPPSI-R for gifted children. The transformation processes between
BSi andDSi, production and remineralization, lead to a continuous cycling between two chemical
forms ofsilicon. Hence, the main objective of this dissertation was to study the motility patterns of
single filaments and the underlying mechanisms. The observation of the single filaments should
provide deeper insight in the behavior on community level. Due to concurrent reoxidation it is not
detectable. Several ecosystem models have been developedfor the North Sea (Radach and Patsch,
1997; Moll, 1998; Skogen and Moll, 2000). So wurden zum ersten Testzeitpunkt (t1) zunachst alle
Kinder untersucht. This has consequences for the strength of the feedback between climatechange
and the marine carbon cycle through changes in stratification and therefore nutrient supply.The
model also is able to reproduce the so-called ’opal-belt’, a region of high deposition of diatomsilica
in the sediment around Antarctica, a feature that is usually not found in fixed-compositionmodels
(not shown).
It also includes many biological statevariables for different organisms. The approach of this study is
to use an ecosystem model for the pelagic, that was developed tosimulate mesocosm experiments
(Schartau et al., 2007), and apply it to the shallow coastal system.The model describes the cycling of
carbon and nitrogen but lacks the consideration of phosphorusand silicon. Schultyp kurzes Intervall
langes Intervall Grundschule 28 28 Hauptschule 1 1 Realschule 15 15 Gymnasium 26 26
Gesamtschule 2 2 Gesamt 72 72 0 2 4 6 8 10 6 7 8 9 10 11 12 13 14 15 16 7 9 43 8 6 6 6 9 9 5 7 9 43
8 6 3 98 78 kurz lang Page 125. Bakken LR (1985) Separation and purification of bacteria from soil.
To illustrate this we calculated Arrhenius plots for each temperature- 58 Page 60. Secular increases
in the Develop-ment Quotients of infants. Ein Kind, mit dem beide Verfahren durchgefuhrt werden,
durfte demnach im HAWIK-III etwas hohere Werte erzielen als im HAWIK-IV. Peterson, J. (1921).
Intelligence and its measurement: A symposium-IX. In winter, chlorophyllconcentrations are almost
zero, although small amounts of chlorophyll can be found in the data. The general explanation for the
similar temperature response in all these different biological processes is that enzyme activity
increases with increasing temperature leading to higher rates up to a critical Topt. In the model,no
iron dependencies appear in the equation for silicon uptake kinetics (Eq. 3.10). Iron shortage
willtherefore not limit silicon uptake directly. The Ea value of 58 kJ mol-1 for Aarhus Bay filaments
adapted to summer conditions was slightly higher than for tropical, cold acclimatized Aarhus Bay
filaments and arctic filaments (49, 50 and 46 kJ 60 Page 62. Below the band, in the sulfidic agar, the
filaments followed random trails with only few reversals. Multi-nutrient modelling of phytoplankton
growth requires knowledge of phytoplankton physiol-ogy and intracellular regulations of different
metabolic processes. Oxygen and brightfield images were taken at intervals ranging from 10-60 min.
Our method is a helpful tool to quickly assess the temperature response of gliding motility in
microorganisms and its application has revealed in recent experiments that Guaymas Basin Beggiatoa
have a psychrophilic response to temperature (H. Although the foodweb structure in the model is
relatively simple, it contains a rathercomplex description of algal physiology that decouples the
uptake of carbon and nitrogen. Jorgensen, 1977, Sayama, 2001, Mussmann, et al., 2003, Preisler, et
al., 2007). Beggiatoa are also widespread in upwelling regions, which are characterized by high
productivity. The assimilationof nitrogen from the water column into the cell requires the transport of
nitrate or ammonia through 45 Page 56. First, detritus is brokenup into dissolved or extracellular
organic carbon and nitrogen, EOC and EON, and second, themore labile EOC and EON is
remineralized into dissolved inorganic carbon and nitrogen, DIC and 19 Page 30.
Phytoplanktoncarbon is additionally reduced by phytoplankton respiration, respphy. London:
Methuen. Jensen, A. R. (1998). The g factor: The science of mental ability. Applied and
Environmental Microbiology 52: 161-168. In the year 2000, phytoplankton carbon is exported into
the NS whereas phytoplankton nitrogen 29 Page 40. The temperature range of gliding correlated
with the climatic origin of the filaments with a high temperature range for tropical, an intermediate
range for temperate, and a low temperature range for arctic filaments. Only at the optimumPOC
uptake rate of the filterfeeders, growth of their own food source, i.e. primary production,
ismaximized. The chemotactic responses available to Beggiatoa in order to form complex community
structures therefore appear to be limited to simple reversals of gliding direction. Dearborn, W. F.
(1921). Intelligence and its measurement: A symposium-XII. In general, phytoplankton
concentrations during the growth period are higher in theWadden Sea than in the North Sea. These
bacteria swim following a relatively straight path which is interrupted by occasional turns. All
population data were recalculated from gram wet weight to volume (cm3) of sediment based on
determined sediment densities.
Postma (1981) proposed the asymmetry of the tidal currents to cause the trapping of particles inthe
sediments of the Wadden Sea, the scour effect. Sediment cores were sampled, generally in the
deeper and central part of the fjords, and the cores were screened for Beggiatoa using the described
approach. The dataset from Helgoland Reede shows much stronger variability 24 Page 35. Fur diesen
Untertest ergeben sich auch dann hohere Werte im HAWIK-IV, wenn diese Test- version zuerst
durchgefuhrt wurde. Such an environmental genomic approach, however, misses the unique
advantage provided by the distinct morphology of Beggiatoa. Phyto-plankton carbon and nitrogen
are reduced by excretion of organic molecules, ?N and ?C, that enterthe pools of dissolved organic
nitrogen and carbon, DON and DOC, respectively. Peterson, J. (1921). Intelligence and its
measurement: A symposium-IX. Fenchel T (2008) Motility of bacteria in sediments. But the
observation of uptake kinetics that linearly increasewith increasing silicon concentrations and the
transition between nonsaturable and saturable siliconuptake kinetics (Thamatrakoln and Hildebrand,
2008) question the role of silicon transporters andwould recommend an alternative pathway to the
transporter-mediated silicon uptake. Thiovulum majus and “Candidatus Ovobacter propellens” cells
perform U-turns when they leave the isopleths of optimal oxygen concentration. Remineralization of
biomass releases all elements that were incorporatedinto biomass, also iron. Also, the penetration
depth of the remotely sensed signal into the water column is notalways known, resulting in relatively
large uncertainties of the remotely sensed data. InAtmospheric Ozone - Proc. 18th Quadrennial
Ozone Symposium L’Aquila,Italy (1996), R. Siegrist, Ed., vol. 127 ofChemical Analysis Series.
Leipzig: Barth. Sternberg, R. J. (1985a). Human intelligence: The model is the message. Sie wird
durch folgenden Regressionskoeffizienten charakterisiert. Complexationby organic ligands prevents
scavenging and keeps dissolved iron bioavailable. Applied and Environmental Microbiology 36: 755-
770. Phytoplankton and zooplankton concentrations during the productive phase of the yearare
usually higher in the SRB than in the NS (Fig. 2.5) and cause a net export of live biomass intothe
NS. Journal of Psychoeducational Assessment, 8, 231-258. A higher exchange efficiency,together
with a stronger influence of Elbe water (with its high winter DIN concentrations) on thewatermass
outside the SRB, may in principle explain the observed DIN peak inside the SRB (Hickel,1980). The
Global Ozone MonitoringExperiment (GOME) makes an important contribution in this field. To
make it a real carbon cycle model, atleast one group of non-silicifying phytoplankton should be
included to improve the simulation ofchlorophyll and nutrient concentrations in ocean regions that
are not dominated by diatoms. Otherimportant functional goups like cyanobacteria and
coccolithophores may be included in the model toaccount for the effects of calcification on total
alkalinity and nitrogen fixation as an additional sourceof reactive nitrogen in nitrogen-limited surface
waters. The high DIN concentrations in the Southern Ocean may be responsible for the high chloro-
phyll concentrations produced by the model. The different variations in the stoichiometric
composition of plankton organisms mentioned aboveindicate that the chemical composition of
seawater can change the stoichiometric signal which is ob-served in plankton biomass. Bern: Huber.
Tewes, U. (1991). Hamburg-Wechsler-Intelligenztest fur Erwachsene - Revision (HAWIE-R). The
model setup (number of cells etc.) was identical to the description above except that the gliding
speed of the filaments was set to 1.4 ?m s-1, which was the gliding speed of the halophile strain.
Butthe release of nutrients by grazers may also support primary productivity. The narrow filaments
had relatively longer cylindrical cells than the wide filaments which had flatter disk-shaped cells.
Woodcock, R. W. (1990). Theoretical foundations of the WJ-R measures of cognitive abilities.
Filaments of these size groups were found to accumulate nitrate in internal vacuoles, often up to
concentrations of several hundred mM (McHatton et al., 1996; Ahmad et al., 1999; Mu?mann et al.,
2003; Preisler et al., 2007). Beggiatoa use nitrate as an alternative electron acceptor when living in
the suboxic zone. The uptake rate in the Michaelis-Menten kinetic (Eq. 3.8), V NC, is regulated in
the same way as the actual photosynthetic rate, ?C. The global distribution of dissolved inorganic
nitrogen, DIN, also shows depletion in surface wa-ters and accumulation towards the lower branch
of the ocean conveyor belt in the deep North Pacific(Fig. 4.2). However, there are major differences
between the global distribution patterns of nitrogenand silicon. Respiration of microphytobenthos
andmacrozoobenthos increases DIC concentrations in the porewaters. The sediment was carefully
placed in a laboratory flume of 3 m length and 0.3 m width without disturbing the layering and kept
in darkness. Diatom growth depletes dissolved silica in the euphotic zone, generating a
verticalconcentration gradient with low DSi concentrations at the ocean surface. Schultyp kurzes
Intervall langes Intervall Grundschule 28 28 Hauptschule 1 1 Realschule 15 15 Gymnasium 26 26
Gesamtschule 2 2 Gesamt 72 72 0 2 4 6 8 10 6 7 8 9 10 11 12 13 14 15 16 7 9 43 8 6 6 6 9 9 5 7 9 43
8 6 3 98 78 kurz lang Page 125. I have implemented the hydrological environ-ment and the pelagic
biogeochemical state variables in both water boxes. Oxygen and sulfide concentration profiles are
indicated with curves 105 Page 107. Image analysis of filaments in the bacterial mat was done in
culture tubes that were not yet densely populated to ensure that individual filaments could be
tracked. Turbulent diffusion and viscosity are parameterized following the scale-dependent
Smagorinsky approach. Beggiatoa occurred throughout the 4 cm deep oxidized zone (Fig. 5) and
two distinct size classes prevailed. DNA is a combination of saccharides, nitrogen rich organic bases
andphosphates. Topt for activities other than growth is commonly far from optimal for the organisms
which then are on the verge of thermal damage, as evident from the steep decline in activity just
above Topt. This pulse occurs after large amounts of benthic detritus have been decomposed
andsediments leave anoxic conditions. After the filament stopped, it pauses for 1-4 s at its stopping
position until it resumes motility (Fig. 10C and D). The resumption of motility is sometimes
accompanied by a sudden jerk into the direction of gliding as if held back by a rubber band that
snaps. Wechsler, D. (1939a). The measurement of adult intelligence. A quadratical loss term
simulates grazing of highertrophic levels on zooplankton. After gentle centrifugation, 0.9 mL of the
supernatant water was collected separately. Filaments in the oxic region above the mat or in the
sulfidic, anoxic region below the mat glided distances longer than the filament length between
reversals. Phytoplanktoncarbon is additionally reduced by phytoplankton respiration, respphy. The
reaction rate (Equ. 1) and its natural logarithm (Equ. 2) can be calculated as RTEa Ae? ?? (Equation
1) RTE A a. Iron is also involved in photosynthesis as it is part ofthe electron transport chain and the
photosystems. Different roles of iron in the cellular metabolismof different elements induce different
regulations of elemental composition under iron-limitation. The determination of the local oxygen
concentration, however, represents a challenge. C) The cells were always rich in light-refracting,
spherical sulfur globules. Several runs are performed withthe salinity model processing a
threedimensional matrix of the unknown model parameters. However, the distribution of elements in
ocean waters, isnot only affected by the formation of biomass. Filaments outside of the mat glided
longer than their own length (Fig. 4A and C). Filaments within the mat glided shorter than their own
length, a reversal pattern that essentially anchored them to the same spot (Fig. 4B). Two particular
filaments were observed as they moved from outside 108 Page 110. The sensitivity towards weak
oxygen gradients provides that Beggiatoa are able to orient even when gliding at an acute angle to
the horizontal (Moller, et al., 1985). A prerequisite seems to be that the oxygen change happens at
low initial oxygen 34 Page 36.
URL Jassby, A. D. and T. Platt. 1976. Mathematical formulation of the relationship between
photosynthe-sis and light for phytoplankton. The water column is assumed to be always 100%
saturated with oxygen due to waves and strongtidal currents. Highervalues of ?Chl lead to very early
phytoplankton growth and even to an anticorrelation with theobserved signal when the bloom period
already starts in winter at very low irradiance levels and endsdue to nutrient depletion and increased
zooplankton grazing. It is not known whether filaments exposed to oxygen concentrations higher
than 10 % air saturation do reverse when they experience an oxygen change of the same magnitude.
This would declare the WaddenSea a net heterotrophic system as it is supported by a net import of
biomass that is remineralized in thesaprophagous food web and cannot be maintained by its primary
producers alone. This procedure improved the statistical representation of the bacterial density in
case these had a patchy distribution. The clonal Beggiatoa culture was populated by an
accompanying organism which is closely related to Pseudovibrio denitrificans (A. Technical report,
NOAA. Levitus, S. and T. Boyer. 1994b. World Ocean Atlas 1994. This uptake mechanism has been
described by a Michaelis-Menten equation(Paasche, 1973b,a; Del Amo and Brzezinski, 1999).
Sulfide is oxidized in two steps: the first step is the oxidation to elemental sulfur. Michaelis-Menten
kinetics are fitted to the data in order to derive maximumsilicon uptake rates and half-saturation
constants for each iron concentration level. However, a comparison of the nitrogensources from
porewater exchange and rivers in the SRB shows that the transport of DIN is directedinto the
sediments in winter (Fig. 2.8) and the major source of DIN is river discharge. This knowledge is
especiallyneeded to make model predictions of future changes of the climate system and its response
to theanthropogenically caused rise in atmospheric CO2 concentrations due to the burning of fossil
fuels. Even though silicon uptake has been shown todecrease under iron limitation too (Leynaert et
al., 2004), iron is not known to be directly involvedin the silicon metabolism of diatoms. Iron is an
important part of the electron transport chain in the 3 Page 14. The reaction rate (Equ. 1) and its
natural logarithm (Equ. 2) can be calculated as RTEa Ae? ?? (Equation 1) RTE A a. Jorgensen BB
(1977) Distribution of colorless sulfur bacteria (Beggiatoa spp.) in a coastal marine sediment. The
Wadden Sea is the adjacient transition zone between the sea and the continent.In the Wadden Sea,
the terrestrial margin has a very gentle slope that falls dry during ebb and isflooded again twice a
day. Diatoms thus grow in the ACC where enough light and DSi are provided. Advective and
diffusive fluxes of dissolved silica,DSi, over four horizontal layers and four latitudinal and vertical
layers are summarized and presented bysolid arrows. The biomass that is builtup in the benthic
compartment may also be exported out of the system by higher trophic levels likeseabirds and fish
that are not considered in this model approach. Gleichung (3.1) ist damit eine skalare Integro-
Differentialgleichung. These coupledbiogeochemical ocean general circulation models, BOGCMs,
can then be used to investigate theregional or global cycles of important biogeochemical elements.
1.3 Study regions This thesis includes four studies that investigate state variable models on different
spatial scales,from phytoplankton batch cultures over regional ecosystems to plankton communities
in the globalocean. Increasing atmospheric concentrations of CO2 increase the ab-sorption of long
wave radiation in the atmosphere and thus induce a heating of the atmosphere (IPCC,2007). Some
Beggiatoa, in particular those of smaller diameters, may thereby have been missed and the counts
thus represent minimum values. IBMs would also require too muchcomputational power and are thus
not suitable for the application in large scale ecosystems or theglobal ocean. Pringsheim EG (1964)
Heterotrophism and the species concept in Beggiatoa. The complete volume of 5 ?L was then used
as template in the following amplification reaction. Discussion The reversal of gliding direction in
entire filaments is apparently achieved by the synchronous reversal of all cells in the filament rather
than by independent reversals by each single cell of the filament. Applied and Environmental
Microbiology 45: 1261-1270. Schultyp kurzes Intervall langes Intervall Grundschule 28 28
Hauptschule 1 1 Realschule 15 15 Gymnasium 26 26 Gesamtschule 2 2 Gesamt 72 72 0 2 4 6 8 10 6
7 8 9 10 11 12 13 14 15 16 7 9 43 8 6 6 6 9 9 5 7 9 43 8 6 3 98 78 kurz lang Page 125.
At temperatures above 5-10?C the filaments glide at a velocity of 2-4 ?m s-1 and effectively migrate
within their preferred 73 Page 75. Ozon wird vonHALOE im thermischen Infrarot (IR ) gemessen.
The narrow 5 ?m Beggiatoa had highest density at 1-2 cm depth while the wider 12 ?m Beggiatoa
had maximum at 2-3 cm depth (Fig. 5A). Due to their much larger individual filament size, the
biomass of the 12 ?m filaments was overall highest (Fig. 5B). The mean areal biomass was 0.167 g
m-2 of which the 5 ?m wide filaments accounted for 40%. 85 Page 87. Thurstone, L. L. (1921).
Intelligence and its measurement: A symposium-X. The numerical proportions that balance chemical
reactions arecalled stoichiometry (Sterner and Elser, 2002). Applied and Environmental
Microbiology 68: 6310-6320. DWD DeutscherWetterDienst ECMWF EuropeanCentre forMedium-
RangeWeatherForecastsInternationale Meteorologische Organisation. Height-resolved ozone
information on a globalscale is required for the detection of changes in the ozone distribution and for
researchinto the underlying chemical and dynamical processes. In most partsof the ocean the
viscosities are much lower. The anticorrelationin these model runs was in fact caused by the high
value for ?Chl that lead to a very early beginningof the diatom bloom and the increasing
anticorrelation is explained by an early breakdown of diatombiomass due to earlier limitation by low
iron concentrations. This knowledge is especiallyneeded to make model predictions of future
changes of the climate system and its response to theanthropogenically caused rise in atmospheric
CO2 concentrations due to the burning of fossil fuels. As a consequence, the slime sheath can
rupture in that region. Applied and Environmental Microbiology 45: 1261-1270. Acknowledgments
We thank Stig Henningsen and John Mortensen for assistance by the field sampling and coring. But
as the growthperiod in the year 2000 is shorter than in the year 2003, less phytoplankton nitrogen is
exported intothe North Sea in the year 2000 than in 2003. High concentrations of oxygen and
sulfide rule in the surrounding environment whereas the mat lives under very low oxygen and sulfide
concentrations. North of the subtropical front, the algae are limited by the supply of nitrogen. This
thesis considers physiological variations in biomass stoichiometry ofone phytoplankton group only
and no variations in the species composition of the plankton commu-nity. 1 Page 12. Gliding speed in
Oscillatoria princes and Flexibacter polymorphus was found to decrease with increasing viscosity of
the medium but showed a linear decrease instead of a curve 62 Page 64. Reversals of gliding
direction happened simultaneously over the entire filament length in the present experiments. The
sediment was mixed silt and fine sand of mostly 20-80 ?m grain size and was light brown and
oxidized to 4 cm depth. At high oxygen flux the filaments accumulate in denser tufts (Moller, et al.,
1985). The tufts are connected by strands of single filaments, giving the mat a web-like appearance.
Such an environmental genomic approach, however, misses the unique advantage provided by the
distinct morphology of Beggiatoa. The zone generally has no detectable sulfide, yet often a high rate
of sulfate reduction. The model by Geideret al. (1998) for variable C:N:Chl ratios in phytoplankton
biomass is chosen as the basis for furthermodel development. Model simulations without riverine
nitrogen discharge lack an important source of DIN in the SRBthat is needed to achieve high DIN
concentrations in winter. Berelson, W. 2002. Particle settling rates increase with depth in the ocean.
They orient in their environment by tactic responses towards chemical stimuli and light. Due to slow
growth of environmental organisms the adaptation does, however, not necessarily track seasonal
temperature variations (Robador et al., 2009). Temperature is an important characteristic of the 51
Page 53. However, some important stoichiometric differences between different types of organisms
exist,mainly caused by different body structures.