Summary of read papers in second week of May

Growth physiology and fate of diatoms in the ocean: a review

Sarthou, et al., 2005

 size is an important factor explaining variations of biogeochemical parameters of diatoms

 variations of elemental ratios can be explained by adaptation of intracellular requirements or storage of Fe, and P, for
instance.
 important loss processes of diatoms pointed out by this synthesis are (i) sinking, as single cells as well as through
aggregation which generally greatly increases sinking rate, (ii) cell autolysis, which can significantly reduce net growth
rates, especially under nutrient limitation when gross growth rates are low, and (iii) grazing by both meso- and micro-
zooplankton.
 Nevertheless, their relatively low surface to volume ratios will need nutrient-rich conditions for growth in
contrast to smaller phytoplankton, such as nano- or pico-plankton, with higher surface to volume ratios, which
allow a more efficient exploi- tation of low nutrient concentrations
 large) diatoms is also observed in man-made perturbations (iron addi- tions) in so-called High-Nutrient, Low-
Chlorophyll (HNLC) regions.
 Values vary by more than one order of magnitude, but they can be related to cell size, which has long been
recognised as an important cause of interspecific variability of Amax, the smallest cells having the highest
growth rates
 umax = 3.4 V-0:13
 The reduction of Amax with increasing cell size implies that small cells have a distinct catalytic advantage over
large ones and should out-compete them
 ncrease in cell size induces a decrease in the capacity of solute influx or efflux on a volume basis, due to a
thicker diffusion boundary layer and a smaller area of membrane lipid and number of transporters allow- ing so
the increase in cell size decreases the effectiveness of increased pigmentation in harvest- ing light, the so-called
‘package effect’
 intracellular concentration Chl-a of diatoms tends to decrease with cell size, resulting in a moderation in the
increase in the package effect with cell size
 shape and the magnitude of the P-E curve reflect the underlying biophysical, biochemical, and metabolic
processes that regulate photosynthesis
 P-E curve can be divided into light-limited, light-saturated and photo- inhibition regions
 Variability in the P-E curve, together with variabil- ity in the Chla:C ratio is used to assess photoacclima- tion,
which involves changes in the macromolecular composition and ultrastructure of the photosynthetic apparatus
 Fe is essential to many metabolic processes that require electron transfer reactions, such as pho- tosynthesis,
respiration and nitrogen assimilation in- ducing that C and N assimilations are directly affected by Fe limitation.
 iron also induces major changes in the photosynthetic characteristics. 80% of the iron required by
phytoplankton is in the photosynthetic electron trans- fer.
 increase in the photosynthetic efficiency was explained by a decrease in light-harvesting pigment content,
reducing the package effect related to the optical properties of a heterogeneous suspension of particles
 reduction in photosynthetic pigment content (chlorosis) accompa- nies Fe limitation
 Therefore, diatoms may dominate phytoplankton community only under conditions of high-nutrient
concentrations. Such con- ditions are found in spring and early summer blooms in open ocean waters. This
implies that at the beginning of the growth season the diatoms should not experience nutrient limitation.
 diatoms can adapt to low nutrient concentrations by reducing their size. Small cells are more proficient than
large cells
 Ethmodiscus has a very high Si:C ratio (0.95 mol/mol), due to a low C content per cell volume
 The largest difference is observed for N:P and Si:P, and could be explained by an intracellular storage of P
 Reduction of size cell is not the only strategy for adaptation to low-iron oceanic conditions (see above). Another
strategy for the adaptation to low Fe concentrations is reducing their Fe requirement for growth
 increase in Si:C and Si:N ratios of diatoms under Fe limitation
 strong impact on Fe limitation on the elemental ratios Si:N and Si:C
 Iron-stressed diatoms should therefore deplete surface waters of orthosilicic acid before nitrate, driving the
system towards Si limitation of the phytoplankton community.
 Si is mainly linked to the growth rate and the duration of the cell wall synthesis phase
 Si assimilation may be indirect- ly affected by Fe limitation through a reduction of growth rate
 Sinking rate of a diatom cell is not only a function of its size and shape but is also strongly dependent on the
physiological state of the cell
 Fe limitation may affect phyto- plankton sinking rates by stressing the energy-producing pathways needed by
the cell to maintain their buoyancy.
 higher silicification of diatoms under Fe-stress may increase the sinking rate via thicker siliceous frus- tules.
 Aggregation processes generally increase sinking rates
 Diatom aggregates are the primary source of marine snow aggregates
 biomass accumulation and concen- tration, cell size and shape, including setae, the abun- dance of chain-
forming species, stickiness, and whether the cells are solitary or chain-forming, and the mechanisms responsible
for particle collision, such as Brownian motion, differential settlement and shear
 One type of cell lysis is the viral lysis, Another type of cell lysis is death due to nutrient stress
 micro- zooplankton are constrained to diets of nanoplankton and strongly suggest that they feed on
phytoplankton larger than themselves

Potential contributions of vertically migrating Rhizosolenia to nutrient cycling and new

production in the open ocean
Richardson et al., 1998

 new primary production due to Rhizosolenia migrations is 17% of new production due to turbulent diffusive fluxes of
nitrate into the euphotic zone and are of the same order of magnitude as new production due to nitrogen fixation in tropical
oceans.
 Large-scale contributions of Rhizosolenia to oceanic new production are limited by their relatively low standing crop.
 Large phytoplankton typically found in these regions include the cyanobacterium Trichodesmium
 large phytoplankters found in open ocean regions have developed mechanisms to cope with the
scarcity of nutrients. Trichodesmium, for example, fixes atmospheric nitrogen, thus avoiding
nitrogen limitation, and uses buoyancy reversals to migrate below the nutricline to acquire
phosphorus
 mat-forming diatom Rhizosolenia and the single-celled diatom Ethmodiscus also use buoyancy
reversals to undergo vertical migrations, allowing them to exploit sources of nutrients in deeper
water
 significantly higher internal nitrate concentrations in ascending cells than in sinking cells, consistent
with the concept of nutrient replenishment by vertical movements between the surface and deep
waters
 If cells are sufficiently abundant, the contributions of Vertical migration may be important for new
production. Further, because nitrate transported to the surface by migrating diatoms may not be
accompanied by stoichiometric equivalents of carbon, new production due to Rhizosolenia
migrations can result in a disproportionately high net removal of carbon from oceanic surface
waters
 Central ocean gyres, where Rhizosolenia is most common, are significant con- tributors to the
export of carbon from the euphotic zone, mostly due to their large geographical area
  The light and nutrient history of Rhizosolenia greatly influences whether cells are positively or
negatively buoyant and the velocity with which cells ascend or sink through the water column

Abundance of the giant diatom Ethmodiscus in the Southwest Atlantic Ocean and Central
Pacific Gyre
Tracy A. Villareal, 1993

 Ethmodiscus Castr. is a widely distributed warm-water diatom noted primarily from

oligotrophic waters
 Ethmodiscus cells are the largest diatoms known, reaching a diameterof2-
3mmandamaximumvolumeof 109p 3

 Pacific Ocean abundance averaged 0.08 cells m−3 at the surface. Atlantic
Ocean/Caribbean Sea abundance ranged from 0.03–4.7 cells m−3 with extensive (150-
fold) spatial and temporal variability.
 viable cells were noted in both upward and downward facing sediment traps at 5400 m in
the Pacific Ocean.
 Extensive sedimentary deposits of Erhmodiscus oozes occur in Pacific, Atlantic and
Indian Ocean sediment
 Several hypotheses have been offered to explain the Occurrence of these deposits, but the
two most widely cited suggest: 1) differential dissolution coupled with resuspension and
deposition, and 2) glacial blooms of Ethmodiscus are responsible for the oozes
 Efhmodiscuscan be positively buoyant
 apparently both rising and sinking, suggests that some components of the dsyphotic zone
hypothesis (deep dwelling populations) may be valid

Buoyancy and growth characteristics of three positively buoyant marine diatoms

J. Keith Moore &Tracy A. Villareal, 1996

 Rhizosolenia - chain- forming marine diatoms which thrive under the low-turbulence,
nutrient-poor conditions typical of the open ocean gyres
 a severe competitive disadvantage in obtaining nutrients relative to small (<5 pm)
phytoplankton due to their lower surface/volume ratio
 Other large phytoplankton in same environment overcome size-related disadvantage by
obtaining nutricline through a buoyancy mediated vertical migration  exploit spatially
uncoupled light and nutrient fields & avoid direct competition with smaller size fraction.
 Positive buoyancy prerequisite for vertical migration.

 vertical migrators directly transport NOf into the upper mixed layer, they cir cumvent the
NO3 trap at the base of the euphotic zone & represent a source of 'new nitrogen' to the
euphotic zone
 large phytoplankton have unusually high C: chlorophyll ratio
 Maximum growth rates ranged from 0.37 to 0.78 divisions d_1 with saturation occurring
between 29 and 164 pmol quanta nr
  Severe growth rate depressions were noted in R. acuminata and R. formosa at irradiance
levels above 50 to 155 pmol quanta m~2 s_
 The percentage of positively buoyant cells was inversely related to light intensity
 In R. formosa both growth rate and tolerance to high light levels decreased substan tially
as cell size decreased
 elevated C: chlorophyll ratios found in buoyant, oceanic phytoplankton are typical of
healthy cells
 carbohydrate ballasting can account for buoyancy changes and that these reserves are
adequate to support dark N03" uptake.
 vertical migration is a property basic to these diatoms’ life history strategy, and, like
multispecies Rhizosolenia mats, solitary Rhizosolenia chains transport new nitrogen to the
euphotic zone in oligotrophic seas.
 inverse relationship between light level and positive buoyancy under nutrient replete
condition

 Supersaturating light  negative buoyancy – Excess carbohydrate at high light  as a

negative correlation between g carbohydrate cell-1 and buoyancy
 R. formosa and R. acuminata also experienced growth inhibition at relatively low light
intensities
 Negatively buoyant mats have very high carbohydrate/protein ratios
 Relationship between buoyancy and light level is a function of size.

Physiological And Optical Properties Of Rhizosoleniania formosa (Bacillariophyceae) in

the Context Of Open-Ocean Vertical Migration’
Tammi L. Richardson,2 Aurea M. Ciotti, John J. Cullen, 1996

 Buoyancy was related to nutrition: Upon N depletion, the per- centage ofpositively
buoyant cells decreased to 4%from 1 1% but reverted to 9% within 12 h of nitrate
readdiction
 Aggregation of chloroplasts was more pro- nounced in N-depleted cells. These are
possibly photoprotective mechanisms that would be a n advantage to N-depleted cells in
surface waters.
 Cells may survive fairly long periods in N-depleted surface waters and will continue to
take up carbon; then they can resume nitrate uptake and revert to positive buoyancy upon
returning to deep, N-rich water,

 Uncoupled uptake of carbon and nitrogen during migrations of Rhizosolenia is a form of

new production that may result in the net removal of carbon from oceanic surface waters.