2ecofisiología de Microalgas

PHYTOPLANKTON ECOPHYSIOLOGY
• Phytoplankton systematics and diversity

• Environmental factors controlling primary
production
• Phytoplankton seasonal succession and
food webs
• Size as ataxonomic alternative for
ecosystem modeling
javier.gilabert@upct.es Technical University of Cartagena

Previous definitions
• PLANKTON, marine and freshwater organisms,

which, because they are nonmotile or because
they are too small or too weak to swim against
the current, exist in a drifting, floating state.
(Encyclopaedia Britannica)
• The plant-like community of plankton is called

phytoplankton, and the animal-like community
is known as zooplankton.

• Autotrophs: They grow on non-organic forms of carbon

and energy using light (or chemical energy).
Phytoplankton are autotrophs – they use CO2 for their
carbon and use sun light for their energy.
• Heterotrophs: organism that obtains its carbon and

energy from organic compounds by feeding on
autotrophs (producers) or of other heterotrophs
(consumers).
• Mixotrophs: organism able to assimilate organic

compounds as carbon sources while using inorganic
compounds as electron donors for energy metabolism.
Some dinoflagellates are mixotrophs.

PHYTOPLANKTON SYSTEMATICS AND DIVERSITY
• Phytoplankton is composed of a large diversity

of single celled micro-organisms (some times
forming colonies) compraising both:
– Prokaryotes: organisms without a cell nucleus

(= karyon), or indeed any other membrane-bound
organelles.
– Eukaryotes: organisms in which the genetic material

is organized into membrane-bound nuclei.

Tree of life
Living
organisms
Eucarya Archea Bacteria
Other Proteo-
Protista Spirochaetes
Eucaryotes bacteria
Dinoflagellates Diatoms Other mega-

Rhodophyta Ciliates Chrysophyta… eucaryotes
Fungi Animalia Plantae
Vascular plants Chlorophyta
Ulvophyceae Chlorophyceae

Taxonomy
• Principal taxonomics groups include:

– Bacillariophyta (diatoms)
– Pyrrhophyta (dinoflagellates)
– Cryptophyta (cryptomonads)
– Chrysophyta (golden-brown algae)
– Euglenophyta (euglenoids)
– Chlorophyta (green Algae)
– Cyanophyta (blue-green algae, cyanobacteria)
– and some others…

Diatoms
Diatoms (Bacillariophyceae)
• Most diverse and
abundant aquatic algae,
dominate phytoplankton
Navicula
of cold, nutrient-rich
water Chaetoceros
• About 5 – 500 μm in Ø
Thalassiosira
pseudonanna
Thalassionema
• Includes both unicellular
and colonial forms
• Common in well mixed
environments
Coscinodiscus
Chaetoceros socialis
http://www.serc.si.edu/labs/phytoplankton/guide/diatoms/diatoms.jsp
Diatoms
• Non-flagellated cells with

external silica cells walls
(frustules) embedded in
pectinaceous matrix
• Frustules are made of
two valves, epitheca and
hypotheca
– Raphe present in some
pennate diatoms enable
gliding movement
– Pores penetrate valves for
entry of gas and nutrients
into the cell

Diatoms
• Two major groups:

– Centric diatoms
(planktonic) are disk-
shaped or cylindrical cells
with radial symmetry
– Pennate diatoms (benthic) Thalassionema
exhibit billateral symmetry

(elongated)
• Pigments: chlorophyll a,
c, carotene, and Fragillaria
xanthophylls

Diatoms
• Cannot swim to maintain

their position in the water
column
• A variety of strategies are
used to prevent sinking:
– Small size
– High surface area/volume Nitzschia pungens
ratio (including spines or
wings) Chaetoceros affinis
– Chain formation
– Actively control their
buoyancy against the
ballast of their shells by
accumulating gas and lipid Chaetoceros coarctatus
storage
– Living in turbulent waters
Asterionella socialis

Diatoms
• Benthic diatoms may

form films on sand
Thalassionema

Dinoflagellates
Dinoflagellates (Pyrrhophytes)
• Most important
phytoplankton besides
diatoms
• They may be plantonic or
benthic (with some
endosymbionst
(zooxanthellae) Thalassionema
• Size range 1 – 2000 μm

• Armored or anarmored
Dinoflagellates
• Two flagella (longitudinal

and transversal) used to
swim vertically
– The longitudinal one
propelles the cell forward
– By undulating motions of
the transversal flagellum
the cell rotates around its
axis while swimming
• Swim up to 500 μm s-1 ~

20 m d-1
• Phototaxis in a number of
species
www.uio.no

Dinoflagellates
• Complex life cycles

• Asexual reproduction
(1d-1)
• Stress can initiate
sexual reproduction
and resting cysts
formation

Dinoflagellates
• Many toxic forms can

cause “red tides”
Geohab
• Toxic species: ca. 60

Dinoflagellates
J. Camp
Strategy ws

Dinoflagellates
• Pigments: Chl a, c,
β-carotene, peridinin
(characteristic accesory
pigment for dynophytes),
gyroxanthin (use to track
toxic blooms of
Gymnodinium breve)
• Abundant in oligotrophic
systems
• Large storage capacity

for phosphorous

Dinoflagellates
• Adapted to low light and

nutrient conditions
• Vertical migration in
numerous dinophytes:
– desdend at night for
nutrient uptake,
– ascend in the morning
for photosynthesis
(enabled by phototactic
response and fast
swimming speeds)

Dinoflagellates
• Extremely vulnerable to
turbulence, so dinophytes
predominate during calm
weather, storms can
destruct large number of
cells

Dinoflagellates
(high) diatoms
Red Tides
dinoflagellates (Thalassiosira)
(low) – Chlorophyll a - (high) large

(Chaetoceros)
(planar) - Form - (rod)
Nutrients
small
(Rhizosolenia)
(Coccolithus)
(Ornithocercus)
(low)
dinoflagellates
(low) (high)

Coccolithophorids
Prymnesiophytes (Coccolithophorids)
• Primarily marine group
• Mostly unicellular
(less than 20 μm), motile (2
flagella)
www.mbari.org
• Highly reflective (water

discoloration can be seen
from satellites)
• Pigments: Chl a, c, β- disc.gsfc.nasa.gov
carotene, many others,

high diversity in accesory
pigments, no pigment
characteristics for all
Cryptophytes
Cryptophytes (cryptomonads)
• Unicellular, naked and motile
with 2 equal length flagella.
Usually flattened dorso-
ventrally.
• With an additional
compartment which contains
nucleid acids (nucleomorph).
Main groups are: Cryptomonas,
• Pigments: chlorophyll a, c, Rhodomonas, Chroomonas)
carotenoids a water-soluble
biliprotein (phycoerythrin or
phycocyanin) as an accessory
antenna-pigment. This
biliprotein is located in the
thylakoid.
Crysophytes
Chrysophytes (golden-brown algae)

• Color is due to dominance of
β-carotene and some
xanthophyll carotenoids (in
addition to chlorophyll a).
• A few species are colonial
• Some (Synura and Uroglena,

for instance) are often major
components when
phosphorous concentration
is low as they have obligate
nutritional requirements for
low P concentration.

Silicoflagellates
Silicoflagellates
• Flagellated cells with
siliceous outer
skeleton
• Small ~ 60 μm
• Common in cold
nutrient-rich-water

Euglenoids
Euglenophyta (Euglenoids)
• Most are unicellular but
lack a cell wall
• 1-3 flagella. Most are

facultatively heterotrophic.
Main groups: Euglena, Phacus.
• Nutrition is supplemented
by the uptake of ammonia
and DON.

Chlorophytes
Chlorophytes (green algae)

• Appearence: mostly bright
green, because chlorophyll is
little masked by accessory
pigments (carotenes, and
some xanthophylls). pediastrium
volvox
• Cell walls consist of inner

cellulose and outer dunaliella closterium cosmarinum
Characteristic genus:
pectinaceous layers. Chlamydomonas, Scenedesmus,
Ulothrix, Pediastrum, Volvox,
Closterium
• Monophyletic group with very

diverse forms that resemble
other forms of algae

Cyanobacteria
Cyanobacteria (Cyanophyceae, blue-green algae)

• Are prokaryotic organisms - they
lack a nucleus and organelles
(chloroplast, mithocondria), have trychodesmium
circular DNA, and have no

chromosomes)
anabaena
• Only phototropic organisms that mycrocistis
perform N2 fixation
• Pigments: Chl a, b, phycobilins, anabaena spirulina
carotenoids
• Unicellular or chain forming
colonies commonly arranged in
filaments – trichomes)
• Asexual reproduction

Cyanobacteria (Synechococcus)
Synechococcus
• A major primary producers on a
global scale in the ocean
• ~ 1 μm diameter
• they acquire major nutrients and

metals at the submicromolar
concentrations found in the
oligotrophic open seas
• their light-harvesting apparatus is

uniquely adapted to the spectral
quality of light in the ocean

Cyanobacteria (Prochlorococcus)
Prochlorophytes
• 0.4 - 0.8 μm diameter
• Prochlorophytes along with Synechococcus,

dominate the photoautotrophic picoplankton in
the world's oceans.
• Instead of the phycobilisomes characteristic of

cyanobacteria, they have divinyl derivatives of
chlorophyll a and b, α-carotene, zeaxanthin,
and a type of phycoerythrin.
• This particular array of pigments enables the

phytoplankter to absorb blue light efficiently at
low light intensities characteristic of the deep
euphotic zone

Index

production
food webs
ecosystem modeling

Index

production:
– Photosynthesis
• Light
• Pigments
– Nutrients uptake
– Temperature
– Grazing

Index

production:
– Photosynthesis
• Light
• Pigments
– Temperature
– Grazing

ENVIRONMENTAL FACTORS CONTROLLING PRIMARY PRODUCTION
PHOTOSYNTHESIS
• Photosynthesis is the process by which

absorbed light energy is used to “split” or oxidize
water, and reduce inorganic carbon dioxide to
organic carbon compounds. Oxygen is produced
as a byproduct of this process.
∗ ∗
~8h ν
CO 2 + 2H 2 O 
→(CH 2O) + H 2O + O2

PHOTOSYNTHESIS
• It takes place in chloroplasts (remind that cyanobacteria

do not have chloroplasts)
• Consists on light and dark reaction

PHOTOSYNTHESIS
Light Reactions
Photosynthetic pigments are organized as “photosystems”
with “antenna” complexes.

PHOTOSYNTHESIS
• Energy absorbed by pigments in the antenna is transferred

to “reaction centers” – specialized chlorophyll a molecules
– where electrons are excited and taken up by the “primary
electron acceptor”

PHOTOSYNTHESIS
Dark Reactions
The Calvin Cycle

uses the products of
the light reactions to
fix carbon dioxide
into organic carbon
compounds.

PHOTOSYNTHESIS
• Photosynthesis require:
– Available solar energy

– Pigments
– Electron transport chains and biochemistry to produce
ATP and ultimately a variety of organic carbon
compounds

LIGHT
c
Light is made of energetic packages λ= (Jules)
called quanta (photons) f
c 1988
The transported energy is given by ε = hf = h = 10 −19
λ λ
and is inversely ralated
to its wavelength
shorter λ  higher E
 larger depth penetration
λ = wavelenght (nm = 10-9 m)

c = light speed (m s-1)
f = frecuency (cycles s-1)
h = Plank’s constant 6,63·10-34
(J s)
LIGHT
Light Extinction
− kλ z
Eλ = E0 λ e
E = light intensity
E0 = incident light
k = extinction coefficient
z = depth

LIGHT
• Extinction coefficient is due to:

– Scattering
– Absorption
• Light is scattered by:

– Suspended materials
– Seston
– Live cells

LIGHT
• Volume Scattering
Function β(θ)
– Radiant intensity (I) in

a given direction from
a volume element
illuminated by a
parallel beam of light,
per unit irradiance (E) 1 dI λ (θ , φ )
on the cross-section of β λ (θ ) ≡ (m-1 sr-1)
the volume, and per Eλ d 3 x
unit volume (d3x).

LIGHT
• Light is absorpted by: - Water

- Dissolved organic matter
0.7
- Detritus
Total - Phytoplankton
0.6
0.5
Absorption (m))
Water
-1
0.4
0.3
Phytoplankton
0.2
0.1
Detritus +
dissolved colored matter
0
400 450 500 550 600 650 700
Wavelength (nm)
PIGMENTS
• Photosynthetic pigments
are responsible for the
absorption of solar
energy.
• All photosynthetic
organisms contain one or
more organic pigments
capable of absorbing
visible radiation, which
will initiate the
photochemical reactions
of photosynthesis.

PIGMENTS
• The three major classes

of pigments are:
– Chlorophylls
– Carotenoids
– Phycobilins.
• Carotenoids and
phycobilins are called
accessory pigments since
the quanta (packets of
light) absorbed by these
pigments can be
transferred to chlorophyll.

PIGMENTS
Chlorophylls
• Contain a porphyrin 'head'
and a phytol 'tail'.
• The polar (water-soluble)
head is made up of a
tetrapyrrole ring and a
magnesium ion complexed
with the nitrogen atoms of the
ring.
• The phytol tail extends into
the lipid layer of the thylakoid
membrane.
– Chlorophyll a - present in all

algae
– Chlorophyll b, c1, c2, d

PIGMENTS
Carotenoids
α-carotene
• Carotenoids contain a conjugated
double bond system of the polyene
type (C-C=C-C=C). β-carotene
• Energy absorbed by carotenoids
may be transferred to chlorophyll a
for photosynthesis; some forms are γ-carotene
photoprotective.
fucoxanthin
– Carotenes:
• α-carotene
• β-carotene
diatoxanthin
– Xanthophylls (oxidated carotenoids)
• Fucoxanthin
• Diatoxanthin zeaxanthin
• Diadinoxanthin
• Peridiniin
• Zeaxanthin … astaxanthin

PIGMENTS

PIGMENTS
Phycobiliproteins
• Linear tetrapyrrols
structurally related to
chlorophyll a but lack the
phytol side chain and
magnesium ion.
• They are water soluble,
unlike chlorophylls and
carotenoids.
• Phycobiliproteins absorb
light in the blue-green
region of the spectrum

PIGMENTS
• Phycobilisome is an
accessory light energy
harvesting structure
mainly composed of
phycobiliproteins (found
in cyanophytes and
cryptophytes )
– Allophycocyanin
– Phycocyanin
– Phycoerythrin
http://www.botany.hawaii.edu/faculty/webb/BOT201/BOT201/Algae
/Bot%20201%20phycobilisome%20hemispherical%20Tsukuba.jpg

PIGMENTS
Light Absorption by pigments
• The “chlorophyll-specific
absorption coefficient”
(a*ph) represents the
absorption - by all active
photosynthetic pigments -
by unit concentration of
chlorophyll a
(m-1 (mg Chl m-3)-1 or
(m2 mg Chl-1)
http://www.iopan.gda.pl/~kaczmar/pracownia/zsinica1.gif
P – E curves
Light – photosynthesis relationship

We call P vs E curves to the curve relating photosynthesis
with light intensity
• Primary production is the

result of photosynthesis
and can be measured in
terms of assimilated carbon
or oxigen released
• Photosynthesis (primary
production) is usually
“normalized” to unit
concentration of chlorophyll
a.

P – E curves
P - E curves
 αE 
• Rectangular P = Pm  
Hyperbola  Pm + αE 
 
 αE 
• Quadratic P = Pm  2 
 P + (αE ) 
2
 m 
• Exponential (
P = Pm 1 − e −αE / Pm )
 αE 
• Hyperbolic tangent P = Pm tanh 
 Pm 

P – E curves
−αEo / Pmax
• At low light, the rate of P = Pmax (1 − e )
photosynthesis is proportional
to the incident (absorbed) light.
The P vs E curve is 1.0
approximately linear with slope
α.
Photosynthesis (g C (g Chl) h )
-1
0.8
-1
– α = Photosynthetic efficiency
0.6
– a*ph is the normalized to

chlorophyll a concentration 0.4
absorption coefficient of
phytoplankton
0.2
– Quantum Yield (Φ) = Photons

Absorbed / Reaction
0.0
(in other words: how many 0 5 10 15 20 25 30
photons are required to yield
some reaction) Irradiance (PAR, µmol m s )-2 -1
α = a Φm *
ph
P – E curves
−αEo / Pmax
P = Pmax (1 − e )
• Pmax is photosynthesis
at the light saturation 5
point (at intermediate
Photosynthesis (g C (g Chl) h )
-1 -1
4
intensities when P-E
curve flattens) 3
2
• The light intensity at the
intersection point 1
between Pmax and α is

Ek 0
0 100 200 300 400 500 600
Irradiance (PAR, µmol m-1 s-1)

P – E curves
• P-E curves changes as 912

a function of growth 12.0 410
200
irradiance.
P (gC gChl-1 h-1 )

10.0 50
9
8.0
PEg
• In a stratified water 6.0
column, the P vs E 4.0
curve changes 2.0

significantly with depth. 0.0
0 500 1000 1500 2000

Irradiance (µmol m s-1 )
-2
• P-E curve at growth
irradiance must be used
in modeling.
PHOTOACCLIMATION
Photoacclimation
• Phytoplankton cells have the ability to
photoacclimate both the intensity and spectral
quality of light.
• Cells adjust their relative amounts of C, Chl and

N contents to maximize growth under the current
environmental conditions.

PHOTOACCLIMATION
At high light cells should :
- synthesize less chlorophyll

(bleached cells)
Carotenoids / Chlorophyll a
0.9
Isochrysis
- protect their photosynthetic 0.8 Nannochloris
apparatus by increasing their 0.7
carotenoids concentrations
relative to chlorophyll a 0.6
0.5
- carotenoids such as β-carotene 0.4

and zeaxanthin do not transfer 1.0 1.5 2.0 2.5
excitation energy to the reaction Log Irradiance (µE m-2 s-1)

center and consequently act to
screen the cell from excess light

PHOTOACCLIMATION
aph* (m2 mg chlorophyll a-1)

Isochrysis
• At low light cells 0.05
0.04
should increase the 0.03
probability of capture 0.02
of photons of light 0.01
(increase of α) by
0.00
0 50 100 150 200 250 300
Irradiance (µE m-2 s-1)

synthesizing more
aph* (m2 mg chlorophyll a-1)

Nannochloris
chlorophyll (greener 0.04
aph*(440)
0.03 aph*(675)
cells)
0.02
0.01
0.00
0 50 100 150 200 250 300
Irradiance (µE m-2 s-1)
Specific absorption coefficient of phytoplankton

a*ph in the red (675 nm) and blue (440 nm) peaks for
Isochrysis taitiana y Nannoclhoris atomus
PHOTOACCLIMATION
• As cells accumulate
chlorophyll, each
chlorophyll molecule
becomes less effective in
light absorption
• This effect is due

primarily to the self-
shading of the
chromophores between
layers of thylakoid
membranes.
Ea ( λ ) / cell = E0 ( λ ) a *( λ ) (Chl / cell )

PHOTOACCLIMATION
• Photoacclimation produces a high variability in

the Carbon / chlorophyll a ratio (10 – 300)
• Carbon / chlorophyll a ratio can be estimated

from light intensity, temperature and nutrients
Chl : C = 0.003 + 0.0154[exp(0.050T )]× (e{− 0.059(I / kH )[1 − exp(− kH )]})× [N / (K N + N )]

(Cloern 1995)

Index

production:
– Photosynthesis
• Light
• Pigments
– Temperature
– Grazing

NUTRIENT UPTAKE
Nutrient uptake
• Synthesis of cell matter requires Nitrogen (N),

Phosphorous (P) and other elements
• Nitrogen is available as nitrate (NO3-) or ammonia

(NH4+), Phosphorous as phosphate (PO4-)
• Phytoplankton acquire nutrients by active uptake

from their water boundary layer

NUTRIENT UPTAKE
Nutrient uptake
• Nitrate is reduced to
nitrite (NO2-) by
nitrate reductase (NR)
• NO2- is reduced to
ammonia (NH4+) in
the chloroplast by
nitrite reductase (NiR)
• NH4+ is incorporated into glutamic acid to form glutamine - via

glutamine synthetase (GS) - then transferred to 2-oxoglutarate –
via 2-oxoglutarate aminotransferase (GOGAT).
NUTRIENT UPTAKE
• Photosynthesis can be
related to nitrogen
concentration
• The total biomass of

any organism will be
determined by the
nutrient present in the
lowest concentration
(Liebig’s law of the
minimum)
• Primary nutrients for

phytoplankton growth
are N and P

NUTRIENT UPTAKE
• Nutrient uptake is limited by the rate at which new

externally-supplied nutrients can diffuse into the
cell boundary layer
• Efficiency of nutrients uptake depends on:
– Diffusive fluxes: how much nutrients reach the cell

surface
– Uptake kinetics: how efficient are the transport
mechanisms

TURBULENCE
Diffusive fluxes

TURBULENCE
• Plankton overcome
diffusion limitation by
actively swimming or by
sinking and turbulence
increases the average
settling velocity of
phytoplankton cells
• Small scale turbulence

affects division rates
Turbulent dissipation rate Sullivan et al. 2003

UPTAKE KINETICS
Uptake Kinetics  S 
V = Vmax  
• The uptake follows the  KS + S 
enzymatic Michaelis-
Menten kinetics 2.5
Specific rate of uptake (h-1)

2.0
– Defined by
• V = uptake rate
1.5
• Vmax = Maximum uptake
rate
• Ks = half-saturation 1.0
constant (substrate
concentration at which 0.5
V = Vmax/2)
0.0
0 1 2 3 4 5 6 7 8 9
Nutrient concentration (µM)

UPTAKE KINETICS
• Phytoplankton growth follows

nutrient uptake. Growth can
be expresses by the Monod
model following the Michaelis-
Menten uptake curve
 S 
µ = µ max  
• μ = rate
 KS + S 
• μmax = maximum growth rate
• Ks = substrate concentration
where growth rate is half of
maximum)
However, phytoplankton cells can store nutrients in internal pools to be

used when they are scarce (luxury uptake)

UPTAKE KINETICS
• Phytoplankton growth is therefore

better described by the Droop
model - based on internal  Qo 
concentration of limiting nutrient µ ' = µ m 1 − 
– μ’ = nutrient limited growth rate  Q
– Q = cell quota
– Qo = minimum cell quota for that
nutrient (cell quota at μ=0)
• Estimation of nutrient-limited
growth rate (μ‘) is more
problematic because of the great
uncertainty about how
fluctuations in nutrient availability
translate into fluctuations in
phytoplankton growth rate.

UPTAKE KINETICS
• Phytoplankton can 2.5

acclimate to chronically Eutrophic waters
low or high nutrient levels Vmax = 2.5 h-1
Specific rate of uptake (h-1)

2.0
Ks = 1.5 µM
adjusting metabolisms for
carbohydrates, lipids, 1.5
pigments and proteins
cell quota (Q) (amount 1.0
per cell)
Oligotrophic waters
0.5 Vmax = 1.5 h-1
• Phytoplankton isolated Ks = 0.25 µM
from oligotrophic 0.0

0 1 2 3 4 5 6 7 8 9
environments have lower Nutrient concentration (µM)
Ks values than
phytoplankton from
eutrophic environments

UPTAKE KINETICS
• photoacclimation also
affects intracellular
quotas:
– Low light:
• More pigments
• Less carbon
• lower C:Chla
– High light:
• Less pigments
• More carbon
• higher C:Chla

NITROGEN UPTAKE
• Most phytoplankton cells preferentially take up

NH4+ over NO3-, and take up nitrate when
ammonium is depleted (some may use organic
sources of nitrogen)

PHOSPHATE UPTAKE
Phosphate uptake
• Alkaline Phosphatase
• Required for ATP synthesis
• Phosphorus occurs mainly as inorganic
phosphate and dissolved organic phosphorus
• Cycles rapidly between phytoplankton, grazers,
and seawater
• Sediments accumulate phosphorus that can be
later released to the water column

STOICHIOMETRY
Stoichiometry
• Chemical composition of
phytoplankton is very responsive
to growth conditions
(e.g. carbohydrates are
accumulated when growth rate is
limited by N)
• The Redfield ratio C:N:P shows

− −
the relationship between 106CO2 + 122 H 2O + 16 NO3 + PO4
dissolved elements in the water
and chemical composition of
phytoplankton growing near their
μmax
(CH 2O)106 + ( NH 3 )16 + H 3 PO4 + 138O2
C:N:Si:P  106:16:16:1

STOICHIOMETRY
0.10
slope = 15.56
Nitrate diffusive flux (mmol m-1d-1)

r2 = 0.98
• In the open ocean the 0.08
P < 0.001
Redfield ratio in the water 0.06
is a consequence of the 0.04
nutrient diffusive fluxes 0.02
through the thermocline 0.00

0.000 0.001 0.002 0.003 0.004 0.005 0.006 0.007
Phosphate diffusive flux (mmol m-2 d-1)
• In coastal waters
Redfield ratio may not
follow

Index

production:
– Photosynthesis
• Light
• Pigments
– Temperature
– Grazing

TEMPERATURE
Temperature
• All chemical processes are
temperature sensitive.
• Rate of metabolic
processes are temperature
rate of process at (T + 10°K)
dependent through the Q10 =
rate of process at T
temperature coefficient, Q10
• The typical temperature

coefficient, Q10, is about 2

TEMPERATURE
P vs E curves (Pmax)
after Valiela 2000

TEMPERATURE
• Growth rate (μ)
Eppley (1972)

Index

production:
– Photosynthesis
• Light
• Pigments
– Temperature
– Grazing

GROWTH
Photsynthesis +
Growth Consumption
Nutrient uptake
µt
N t = N 0e
Doubling time = ln2/μ

GRAZING
Grazing
• Phytoplankton is Oithona Daphnia
eaten by zooplankton
(e.g. copepods in
marine waters, and
cladocerans in
freshwater)
• Copepods have
external mouthparts

GRAZING
Cilliates

GRAZING
Ivlev model
− kp
r = R(1 − e )
r = amount of food ingested
R = maximum ratio taken
p = prey density
MaAllister model
I = I m (1 − e − A( P − P0 ) )
I = rate of ingestion of phytoplankton
Im = maximum rate of ingestion
P = mean phytoplankton concentration during grazing period
P0 = minimum phytoplankton concentration required to induce filtering
A = constant defining the rate of change of ingestion with food concentration

GRAZING
Grazing is also a
temperature-dependent
processes

Index

production
food webs
ecosystem modeling

PHYTOPLANKTON SEASONAL SUCCESSION
Varying environmental
conditions produce
seasonal variations on
phytoplankton
Successional pattern (Margalef):
stage I: fast-growing, small-celled diatoms

stage II: larger diatoms with lower growth rates
stage III: large dinoflagellates with still lower
growth rates.

Mar Menor lagoon

Phytoplankton
2000
Diatoms (90%) Diatoms (80%)
Dinoflagellates (30%)
Density (cells · mL-1)
Rhodomonas (40%)
Euglenals (2%)
1500 Pyramimonas (8%)
Diatoms (20%)
Rhodomonas spp. (85%)
Cyclotella spp. (70%) Rhodomonas (70%)

Small Dinofl. (15%)
1000
Small Diatoms (20%)
Small Dinofl. (20%)
Rhodomonas (40%)
Diatoms (15%)
500
0
J F M A M J J A S O N D J
Time (months) Gilabert, 2001

Diatoms
2000
Nitzschia cf. closterium
1500
1000 Cyclotella
Chaetoceros
500
0
Dinoflagellates
400
Ceratium furca
350
200
100
0

ZOOPLANKTON
Mar Menor lagoon

Zooplankton
2000
Nauplii + Copepods (95%)
Copepods (40%) Tintinnids (83%)

Copepods (3%)
Density (Ind · L-1)
Appendicularia (2%)
1500 Veliger larvae (40%) Veliger larvae (7%)
Copepods (80%)
Copepods (80%)
Tintinnids (80%)
Copepods (10%)
1000 Eggs (5%)
Copepods (35%) Larvae (5%)
Nauplii (55%)
Appendicularia (2%) Copepods (40%)
Veliger larvae (20%)
500
0
Time (months)
FOOD WEBS
How can we model phyplankton succession and food webs?

FOOD WEBS
Species change with trophic status

A
Gamito et al 2005, CRC press chapter 5

FOOD WEBS
Sieburth, 1974
Fenchel, 1974

FOOD WEBS
Gamito et al 2005, CRC press chapter 5

FOOD WEBS

FOOD WEBS

OUTLINE

production
• Phytoplankton seasonal succession
• Planktonic food webs
ecosystem modeling

SIZE
ALLOMETRY
Allometry
• The study of size and its consequences (Gould, 1966)
• Body size has an influence on organisms on

physiological, ecological and evolutionary scales
• The consequences of size are therefore expressed as

power functions (exponents)
R = aM b
R = Metabolic rate
a = constant
M = body mass
b = describes effect of body mass on variable R
SIZE
ALLOMETRY
Some basic allometric

relationships are:
Surface area : Volume

• Metabolic rate vs Volume
– In an sphere
• Surface area = 4 π r2
• Volume = 4/3 π r3
• Surface area to Volume ratio
– SA/V = 3/r ∝ V-1/3 length
• Metabolic rate per unit

volume can be related to
SA/V:
– SA/V = 3/r ∝V-1/3
• Metabolic rate per
individual:
– SA/V ⋅V ∝ V-1/3 ⋅V1 ∝ V2/3
SIZE
Allometry has received much attention in physiology, but only recently is

taking importance in ecology
Max Kleiber

SIZE
Allometrical basis of phytoplankton ecophysiology
Surface Area to Volume

ratio in phytoplankton
Reynolds, 1984

SIZE
METABOLIC PROCESSES
• Ecophysiological processes depending on cell

surface should be size-dependent
– Light absorption
• Pmax vs size
• a*ph vs size
– Nutrient uptake
• Kinetics (Ks vs size)
• Diffusion
– Sinking
– Grazing
• Predator and Prey size

SIZE
METABOLIC PROCESSES
Light absorption
• Pmax vs size
Changes in exponents are associated with growth

under resource-limiting versus saturating conditions

SIZE
METABOLIC PROCESSES
Light absorption
• chlorophyll-specific absorption coefficient (a*ph)

SIZE
METABOLIC PROCESSES
• Nutrient uptake
– Ks vs size Size-dependence of Ks suggests that small celled
phytoplankton are better adapted to grow in
oligotrophic conditions than larger ones
Malone, 1980
SIZE
METABOLIC PROCESSES
• Nutrient uptake
– Vmax vs size
Hein et al. 1996

SIZE
METABOLIC PROCESSES
• Nutrient uptake
– Diffusion
J = 4πrDS
J = Supply of nutrient to the cell

r = radius of the cell
D = molecular diffusion coefficient
S = ambient concentration
Chisholm, 1992
Steady state diffusion for a sphere with zero boundary conditions
SINKING
• Sinking
1 ( ρ1 − ρ 2 ) 2
– Stokes law v= g d
18 µ
v = sinking speed
Small particles remain for long d = diameter
time in the water column ρ1 = particle density
ρ2 = water density
μ = water viscosity
g = gravity

SIZE
METABOLIC PROCESSES
• Growth rates are also size-dependent:

– Maximum growth rate

SIZE
METABOLIC PROCESSES
• Growth rates are also size-dependent:

– Turnover rates (P:B)
– Density
Sandow 2004

SIZE
PRIMARY PRODUCTION SIZE-SPECTRA
Thank you for your attention
I have used numerous sources for concepts and figures, references for this sources are available upon
request by e-mail

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PHYTOPLANKTON ECOPHYSIOLOGY

• Phytoplankton systematics and diversity

javier.gilabert@upct.es Technical University of Cartagena

• PLANKTON, marine and freshwater organisms,

• The plant-like community of plankton is called

javier.gilabert@upct.es Technical University of Cartagena

• Autotrophs: They grow on non-organic forms of carbon

• Heterotrophs: organism that obtains its carbon and

• Mixotrophs: organism able to assimilate organic

javier.gilabert@upct.es Technical University of Cartagena

• Phytoplankton is composed of a large diversity

– Prokaryotes: organisms without a cell nucleus

– Eukaryotes: organisms in which the genetic material

javier.gilabert@upct.es Technical University of Cartagena

Eucarya Archea Bacteria

Dinoflagellates Diatoms Other mega-

Fungi Animalia Plantae

Vascular plants Chlorophyta

javier.gilabert@upct.es Technical University of Cartagena

• Principal taxonomics groups include:

javier.gilabert@upct.es Technical University of Cartagena

• Non-flagellated cells with

javier.gilabert@upct.es Technical University of Cartagena

• Two major groups:

exhibit billateral symmetry

javier.gilabert@upct.es Technical University of Cartagena

• Cannot swim to maintain

javier.gilabert@upct.es Technical University of Cartagena

• Benthic diatoms may

javier.gilabert@upct.es Technical University of Cartagena

• Size range 1 – 2000 μm

• Two flagella (longitudinal

• Swim up to 500 μm s-1 ~

javier.gilabert@upct.es Technical University of Cartagena

• Complex life cycles

javier.gilabert@upct.es Technical University of Cartagena

• Many toxic forms can

• Toxic species: ca. 60

javier.gilabert@upct.es Technical University of Cartagena

javier.gilabert@upct.es Technical University of Cartagena

• Large storage capacity

javier.gilabert@upct.es Technical University of Cartagena

• Adapted to low light and

javier.gilabert@upct.es Technical University of Cartagena

javier.gilabert@upct.es Technical University of Cartagena

(low) – Chlorophyll a - (high) large

javier.gilabert@upct.es Technical University of Cartagena

• Highly reflective (water

• Pigments: Chl a, c, β- disc.gsfc.nasa.gov

carotene, many others,

Chrysophytes (golden-brown algae)

• A few species are colonial

• Some (Synura and Uroglena,

javier.gilabert@upct.es Technical University of Cartagena

javier.gilabert@upct.es Technical University of Cartagena

• 1-3 flagella. Most are

javier.gilabert@upct.es Technical University of Cartagena

Chlorophytes (green algae)

• Cell walls consist of inner

• Monophyletic group with very

javier.gilabert@upct.es Technical University of Cartagena

Cyanobacteria (Cyanophyceae, blue-green algae)