BIOLOGICAL OCEANOGRAPHY

3 From shallow to deep sea environments - Distributions of the classical, major

marine systems (kelp beds, mangroves, seagrasses, coral reefs and sediments)
4, 5 How to sample the oceans. Definition of basic terms (plankton, pelagic etc.).
Introduction to the major groups of organisms, particularly phytoplankton, in the
marine food web. The Microbial Loop. Primary production. What limits
photosynthesis in the sea?
6 Photosynthetic picoplankton - Discovery, physiology, adaptation to light climate,
seasonal cycles, differences in pigment composition between the genera
Synechococcus and Prochlorococcus
7 Biogeochemical cycles. The role of micro-organisms in the cycling of carbon,
nitrogen, phosphorus and sulphur
8 Nitrogen fixation in the sea; Microbiology of hydrothermal vent communities and
adaptation to living at depth
9 Marine microbial interactions - From single cells to streamlined genomes and
microbial interdependence; ecological theories (red vs black queen); positive and
negative interactions, viral lysis and grazing.
10 Molecular approaches to assessing genetic diversity in the marine environment;
the 'great plate count anomaly'; 16S rRNA analysis; PCR approaches.
11 Anthropogenic effects on the oceans – increasing atmospheric CO 2 and the
buffering effect of the oceans, ocean acidification, eutrophication and harmful
blooms.
FOOD CHAINS, ENERGY TRANSFER & THE MICROBIAL LOOP
 
Why we need to understand carbon flux pathways:
 
i) in order to predict whether the oceans will respond to increasing
atmospheric CO2 by acting as a net CO2 sink.
 
ii) biodiversity & fisheries management also requires predicting
spatial-temporal patterns of carbon fluxes into various biomass &
species pools.
 
 
Organic matter synthesized by phytoplankton can enter 3 main
pathways:-
 
i) it can be eaten by protozoa & metazoa (zooplankton)

ii) it can be utilised by bacteria

iii) it can aggregate & sediment

Classical Food Web :
phytoplankton  zooplankton  fish (N.B - no bacteria)
Now know that metabolism of bacteria & protozoa strongly influence
carbon fluxes
Microbial Loop : term first used by Azam (1983) incorporating the
role of bacteria & planktonic protozoans in marine food webs.
Bacterial biomass roughly = phytoplankton biomass
Bacterial C demand is often equivalent to about half of the primary
production
Thus major fluxes of organic matter occur via the route:
phytoplankton  DOC  bacteria  protozoa.
phytoplankton DOC bacteria

protozoa
Simplified schematic
diagram of the
‘Microbial Loop’ microzooplankton

zooplankton
Coupling of the pelagic grazing food chain (phytoplankton to fish) and the
Microbial Loop (bacteria & protozoans). Dashed lined show the release of
DOC as metabolic by-products used as a source of C by heterotrophic
bacteria. The bacteria are consumed by protozoans which are in turn
eaten by larger zooplankton.
The Microbial Loop is fuelled by dissolved organic matter:
 
This is derived from
 
i) excretion of DOC by phytoplankton & cell lysis,
 
ii) ‘sloppy feeding’ by zooplankton which can’t feed efficiently on small
particles, and release DOC
 
These findings have changed the way that oceanographer’s think of the
‘ocean’s metabolism’, as well as ideas of food web structure & carbon
flux pathways
 
So, in the microbial loop a significant proportion of primary production
is channelled through detrital material & heterotrophic micro-
organisms.
 
This reduces ecosystem efficiency since there is only a ~10% efficiency
of energy transfer between trophic levels (much of the chemical energy
incorporated into organic compounds is converted to heat energy & is
dissipated in respiration).
PRIMARY PRODUCTION
 
 Fixation of CO2 to produce biomass
 
 greatest part is photosynthetic by phytoplankton
 
 recent discovery that chemoautotrophic bacteria can support animal
communities at hydrothermal vents
 
 phytoplankton restricted to the surface mixed layer - the euphotic
zone or depth is the depth that photosynthetically available/active
radiation (PAR) 400 -700 nm penetrates
 
 compensation depth is the depth in the water column where irradiance
supports a rate of photosynthesis which balances respiration
 
 primary production includes DOC excreted by phytoplankton which is
available to, & utilised by, bacteria
METHODS OF MEASURING PHYTOPLANKTON BIOMASS
a) Pigments:
 
 chlorophyll a (chl a) is the major light harvesting pigment - present in all
photosynthetic organisms it is thus a good indicator of phytoplankton biomass
 
 chl a is easily measured by fluorometry in which the sample is illuminated
with blue light, chl a emits red light as fluorescence
 
 fluorometers are robust & can be submerged to depths of 1000 m or more &
can be used for continuous recording, producing much information
 
 other pigments (accessory pigments) can be used as taxonomic markers
specific for particular phytoplankton groups e.g. fucoxanthin in diatoms,
phycoerythrin in cyanobacteria; the disadvantage of these is the method of
analysis - require to use high pressure liquid chromatography (HPLC) which can
take as long as 30 min per sample. Also, phycobiliproteins are water soluble
 
b) Microscopy
 
c) Electronic particle counter (e.g. Coulter counter)
 
d) Flow cytometry & image analysis
METHODS OF MEASURING PHYTOPLANKTON PRODUCTION
 
a) 14C incorporation (incubations following incorporation into biomass)
 
b) oxygen production in photosynthesis (need to account for respiration)
 
Photosynthesis : CO2 + 2H2O  CH2O + O2 + H2O
 
14
C Method - measures a rate
 
i) NaH14CO3- is added as a tracer to an in vitro incubation a) in the light & b) in
the dark
 
ii) Amount of radioactivity taken up per unit time is then measured on the
filtered phytoplankton.
 
iii) Primary production (in mg C m-3 h-1) is calculated from:
  R = total radioactivity added to the sample
rate of production = (RL - RD) x W N = number of hours of incubation
R x N  RL = counts in the light sample
RD = counts in the dark sample
W = estimate of the total HCO3- content
of seawater
LIMITS TO PHYTOPLANKTON PRODUCTION
 
i) Physical processes
 
Light
Temperature
Vertical Mixing
Advection
Stratification
 
ii) Chemical processes
 
nutrient supply : N P Si, trace metals, CO2
 
iii) Grazing by microzooplankton, copepods, jellyfish etc
 
iv) Infection e.g. by viruses
LIGHT IN THE SEA
 
 water is relatively transparent to solar radiation
 
 of the sunlight penetrating the sea surface about 50% is composed of
wavelengths longer than 780 nm. This infrared radiation is quickly
absorbed and converted to heat in the upper few metres
 
 ultraviolet radiation (<380 nm) forms only a small fraction of the total
radiation & is rapidly scattered and absorbed, except in very clear ocean
waters
 
 remaining 50% of the radiation comprises the visible spectrum,
wavelengths of 400 -700 nm that penetrate deeper in the sea
 
 in oceanic regions blue-green light penetrates the deepest
 
 in coastal regions green light penetrates the deepest
 

 there are global variations & seasonal effects in the amount of sunlight
whilst also phytoplankton are often being mixed from areas of bright
light to areas of dim/little light
Variation in light intensity

Variation in light quality

SEASONAL CYCLE OF PHYTOPLANKTON PRODUCTION IN
TEMPERATE WATERS
Winter
  Low production rates because the surface mixed layer is deep since there is no
stratification
· phytoplankton undergo deep mixing and can be ‘mixed out’ of the euphotic zone.
Nutrients are high, but there is insufficient light to be utilised
Spring
 surface warming results in suppression of deep vertical mixing. The thermocline
begins to form. This small amount of heating stabilises the water column so
there is no deep mixing.
· phytoplankton are still mixed from high to low light but are able to
photosynthesise for a greater proportion of the day.
· growth is maximised, nutrients are still plentiful & a high biomass develops (the
spring bloom)
· diatoms dominate
Late Spring
 surface heating continues & the thermocline becomes stronger
 high biomass of phytoplankton with concomitant utilisation of N, P & silicate
· diatom growth stops & senescent cells begin to sink
· zooplankton having longer generation times than the phytoplankton, hence there
is a lag between the increase in phytoplankton & the increase in grazers which
can exploit the food resource
Late Spring (cont.)
· as grazing activity increases there is an increase in zooplankton activity
(respiration) and increased excretion of ammonia (whereas nitrate can’t
penetrate through the thermocline from the deep water below)
· the regenerated nutrients are utilised by phytoplankton, though primary
production is maintained at a lower level than during the spring bloom
· silicate recycling is much slower than N or P, hence there is insufficient silicate
for diatom growth & microflagellates replace the diatoms as the dominant
phytoplankton
· with lower photosynthetic biomass in the water column, light can penetrate as
far as the thermocline, where nutrient concentrations are relatively high. A
deep chlorophyll maximum results
Late summer
 Increased grazing activity results in more re-generation of nutrients & there
can be a late summer bloom. Dinoflagellates may sometimes dominate giving rise
to ‘red tides’
 
Late Autumn
 more heat is lost from the surface mixed layer than is gained
 the temperature difference between surface & deep waters is reduced & the
thermocline is eroded
 phytoplankton are mixed below the euphotic zone & production declines to
winter values