BIOGEOCHEMICAL CYCLES

• Chemical elements including all essential elements of protoplasm circulate in the

biosphere in characteristic paths from environment to organism and back to
environment.
• Paths → Biogeochemical Cycles
• Elements & inorganic compounds essential to life,
→ Nutrient Cycles
• Energy Flow
heat heat
↑ ↑
Sunlight → Producers → Consumers
↓
heat - one way process

• Biogeochemical Cycles
• no starting point
• each segment is dependent on the one preceding it
• emphasizes global interconnectedness of organisms (bio) and the rest of
the earth (geo)
• Odum • 2 pools • reservoir • large, slow-moving, non-biological component
• exchange or cycling pool • smaller but more active portion
exchanging between organism & environment
• 2 groups of biogeochemical cycles
• Gaseous types • reservoir is in the atmosphere or hydrosphere (C,N,O)
• Sedimentary types • reservoir is in the earth’s crust (P, S, Fe etc)
 Algae and Biogeochemical Cycling
Biogeochemical cycling: movement and exchanging of both matter and energy on
earth  atmosphere, hydrosphere, lithosphere and the biosphere

Algae provide biogechemical services.

Cyanobacteria,
through oxygenic
photosynthesis
transformed the
atmosphere

Earth’s initial
Present
atmosphere:
composition: 78%
80% N2, 10% CO/CO2,
N2, 21% O2,
10% H2 (by volume) 
0.036% CO2
no free O2
Marine & Freshwater Algae like Eustigmatophyceae, Dinophyceae, Chlorophyceae
 Organic matter  fossilized hydrocarbons  fossil fuel

Calcareous algae like coccolithophorids, coralline algae  calcium carbonate

source  bone grafts; maerl fertiliser

Diatoms  diatomaceous earth (diatomite)  filter aids

Algae : 99% of biomass comprises C, O, H, N, S, P, Ca, K, Na, Cl, Mg, Fe, Si

• Algae die and return the organic materials back to inorganic forms of elements 
Mineralisation

• Takes place in water column and bottom of pond/lake or sea/ocean

• Recycling is rapid – in an euphotic zone

Or slow – eg. refractory compounds that accumulate on seabed

• In water column with O2  heterotrophic bacteria carry out the oxidative

degradation
•  CO2 and nutrients released for reuse

• Nutrients that may be limiting  nitrate (NO3−), iron (bioavailable Fe), phosphate
(PO43−), and dissolved silicon (Si(OH)4) are most often found in concentration
well below the half-saturation levels required for the maximum phytoplankton
growth

• Algae are important for the biogeochemical cycling of the chemical elements they
uptake, assimilate, and produce such as carbon, oxygen, nitrogen, phosphorus,
silicon, and sulfur.

Basanti L & Gualtieri P. (2014). Algae.

Anatomy, Biochemistry and Biotechnology
(2nd Edition). CRC Press.
Liebig’s “Law of the Minimum” :

Growth is controlled not by the total of nutrients available but by the nutrient available
in the smallest quantity with respect to the requirements of the plant

“When other factors such as light & temperature are favourable, the nutrient available
in the smallest quantity, is limiting to the growth of the seaweed”

Nutrient-limited growth is usually modeled with a Monod (or Michaelis–Menten)

equation:
m= mmax [LN]
________
[LN] + Km

where μ is the specific growth rate of population as a function of [LN],

[LN] is the concentration of limiting nutrient,
μmax is the maximum population growth rate (at “optimal” conditions), and
Km is the Monod coefficient, also called the half-saturation coefficient because it
corresponds to the concentration at which μ is one-half of its maximum.

When the concentration of limiting nutrient [LN] equals Km, the population
growth rate is μmax/2.
As [LN] increases, μ increases and so the algal population (number of cells)
increases.

Beyond a certain [LN], μ tends asymptotically to its maximum (μmax), and

the population tends to its maximum yield.

If this concentration is not maintained, primary productivity returns to a level

comparable to that prior to the nutrient enrichment rapidly  seasonal
blooming.

But when there is a continuous excess supply of one or more limiting

nutrients, which leads to intense and prolonged algal blooms throughout the
year  giving a constant maximum algal growth rate (μmax)  leads to a
continuous primary production.

 When this occurs, we refer to it as eutrophication.

 Limiting nutrients : N, P, Si
N becomes limiting in high salinities and low flow.
P becomes limiting in freshwaters, intermediate salinities

These nutrients are present in algal cells in a species-specific structural ratio, the
so-called Redfield ratio, which determines the nutrient requirement of the species

Redfield Ratio depends on the conditions under which species grow and
compete.

Phosphorus limitation:

• In phosphorus limitation: growth is limited; limitation of nucleic acid synthesis  affect

photosynthetic energy conversion by reducing the rate of synthesis of proteins in the photosynthetic
apparatus [at the level of genome replication or at the level of RNA synthesis (a form of
transcriptional control) ]  affect cell metabolism and oxidative stress

• Reduced rate of synthesis and regeneration of substrates in the Calvin–Benson cycle, thereby
reducing the rate of light utilization for carbon fixation.

• Decrease in membrane phospholipids

• Inability to produce nucleic acids under P limitation limit cell division, leading to an increased cell
volume.
N limitation:

• directly influences supply of amino acids  limits the translation of mRNA and
hence reduces the rate of protein synthesis.

• efficiency of PSII decreases, consequence of thermal dissipation of absorbed

excitation energy in the pigment bed  due to a decrease in the number of
PSII reaction centers relative to the antennae.

• As nitrogen limitation leads to a reduction of growth and photosynthetic rates, it

also leads to a reduction in respiratory rates. The relationship between the
specific growth rate and specific respiration rate is linear with a positive
intercept at zero growth, which is termed “maintenance respiration.”

• The molecular basis of the alterations is unclear; however, the demands for
carbon skeletons and ATP, two of the major products of the respiratory
pathways, are markedly reduced if protein synthesis is depressed.
Silicon limitation:

• Specifically for diatoms

• Silicic acid uptake, silica (SiO2) frustule formation, and the cell division
cycle are all tightly linked

• Under silica limitation, the diatom cell cycle predominantly stops at the
G2 phase, before the completion of cell division  an inhibition of cell
division linked to an inability to synthesize new cell-wall material under
silicon limitation can lead to an increase in the volume per cell.

• This increase could also be partly explained by the formation of

auxospores with a larger cell diameter.
Sulphur

• Sulfur is an essential element for autotrophs and heterotrophs.

• Important in the structure and function of proteins.

• Three amino acids found in almost all proteins (cysteine, cystine, and
methionine) contain carbon-bounded sulfur.

• Sulfur is also found in sulfolipids, some vitamins, sulfate esters, and a variety of
other compounds.

• Sulfate is taken up into cell by active transport mechanism  converted to

PAPS (3′-phosphoadenosine-5′-phosphosulfate)  transferred to a thiol carrier
(a molecule with a –SH group) and reduced to the −2 oxidation state.

• In sulphur assimilation, the sulfur remains attached to a carrier during the

reduction sequence. In a final step, the carrier-bound sulfide reacts with O-
acetyl-serine to form cysteine.

• Cysteine serves as the starting compound for the biosynthesis of all other sulfur
metabolites, especially the other sulfur-containing amino acids homocysteine
and methionine.
 SHET 2105 L3.1 Nutrition
Nutrient requirements:
•56 elements found in marine algae
•C, H, O. K, N, S, P. Ca, Mg > 1mg/g DW

a) Macronutrients:
1. Carbon  in the form of CO2, HCO-3, CO-3, organic compound
2. Nitrogen  nitrate, nitrite, ammonium, organic nitrogen
control growth; nitrate can be stored in vacoules eg. Valonia
3. Phosphorus  orthophosphate ion (1-3 μM)
 in seawater at pH 8.0 (20°C): 87% of phosphate is in the form of
HPO42-; 12% PO43-; 1% H2PO4-
 store polyphosphates
4. Sulphur  sulpholipid, sulphated polysaccharide
5. Calcium  in cellular membrane; skeletal structure; coral
6. Magnesium  component of chlorophyll; involved in aggregation of ribosomes
into functional units
7. Potassium  activator for several enzyme systems

b) Inorganic Micronutrients:
catalyst in metabolic reactions; osmoregulation
1. Zinc  Zn deficiency  decrease in growth; decrease in macromolecular
protein; decrease in chlorophyll, phycobiliprotein
2. Halogen  Iodine growth, morphogenesis, reproduction
 Bromine – growth
 Chlorine – photosynthesis
3. Silicon  important for diatoms
4. Cobalt  for synthesis of vitamin B12
5. Iron  structural component; cofactor in enzyme reactions
6. Boron  embryo growth
7. Arsenic  involved in phosphorus metabolism
8. Copper  toxic if >100μgL-1
9. Manganese  important in photosynthesis

c) Vitamins & plant growth regulators:

•Supplied by microflora  microalgae, bacteria, fungi (contaminants in seaweed
culture)
•Thiamine, vitamin B12

Effect of Nutrient Deficiency:

•Decrease in photosynthetic pigments
•Store C-storage compounds
•Decrease in protein & nucleic acids
•Increase in nutrients  vegetative growth & asexual reproduction
•Decrease in nutrients (N)  gametes produced
Nutrient Absorption:
Mechanisms:
•Nutrient enters cell as cation &
anion
•Passive diffusion in free space
•Ion-exchange system 
adsorption to cell wall (which
contains polysaccharide,
mucoprotein)
•Carrier system  accumulation
against the electrochemical
gradient
•Active transport  requires
energy
•Influenced by : light, temperature,
water movement, ion content
Requirements for developing nutrient recipes for
algal cultivation as follows:
 Nutritional modes
Amphitrophy
Autotrophy Heterotrophy
Mixotrophy

Photoautotrophy
Photoheterotrophy

Auxotrophy

Obligate
Photoautotrophy
Amphitrophy: can use either autotrophy
or heterotrophy
Auxotrophy: cannot synthesis specific
organic molecules
1. Autotrophy
Cells obtain their energy through absorption of light energy for the reduction of
carbon dioxide by the oxidation of water with the release of oxygen.
Require small amounts of essential organic compounds like vitamins, amino-acids

2. Photoautotrophy
Autotrophic cells that require only inorganic mineral ions
Obligate photoautotrophs cannot grow in dark

3. Heterotrophy
Cells obtain their energy needs from organic compounds produced by other
organisms

4. Photoheterophy
Cells require light as an energy source to use the organic compounds as nutrients.
The organic compounds may satisfy the energy requirements of the algae.

5. Mixotrophy
Like autotrophy but where organic carbon and carbon dioxide are required

6. Amphitrophy
Like heterotrophy but where organic carbon and carbon dioxide are required
Carbon Metabolism
1. In photoautotrophic organisms: light energy

NADPH + ATP
2

CO carbon compound
2

2. Sources of inorganic carbon: dissolved CO , H CO , HCO -1, CO -1

2 2 3 3 3
: content depends on pH, temperature, salinity
Carbonate equilibrium:
pk=6.0 pK= 9.1
CO + H O ↔ H CO ↔ HCO -1 + H+ ↔ CO 2- + H+ ↔ HCO -1 ↔ CO
2 2 2 3 3 3 3 2

+ OH-1

-1 is the dominant form (90%)

• pH of seawater is between 7.8-8.2; at this pH, HCO
3
 carbonic anhydrase enzyme

3. HCO -1 → CO for ‘primary’ carbon fixation enzyme of

3 2
photosynthesis, ribulose-1, 5-biphosphate carboxylase
4. Many species of algae can use organic carbon (heterotrophic, mixotrophic): Chlorella
can use glucose, acetate, leucine

HCO -1 CO
3 2 CO HCO -1
2 3

eg. Chlorella

7. Intertidal seaweeds can use CO2 when exposed to air during low tide

8. Diffusion rates of carbon dioxide from atmosphere into water can only support
algal productivities of 10 g (DW) m-2 d-1

9. During photosynthetic carbon fixation, pH rises to as high as 11 if no

additional CO2 is supplied
Nitrogen
• Nitrogen content of biomass can range from 1 – 10%
• During N limitation, discolouration occurs (decrease in chlorophyll and
increase in carotenoids); accumulation of polysaccharides & oils (PUFAs)
• Ammonia-nitrogen is the preferred source
• When ammonia is utilised, pH can drop – due to release of H+ ions
 Can use this to control predators

• At high pH, ammonia can be volatilised & lost

• Cyanobacteria can use elemental N by reduction to NH4+; catalysed by
enzyme nitrogenase

Phosphorus
• Essential for growth; energy transfer, biosynthesis of nucleic acids, DNA
• Preferred form is ortho-phosphate (PO42-) ; uptake is energy-dependent
• Biomass contains <1% but important because bound to other ions (eg. CO32-
, iron) resulting in its precipitation & unavailability
• Algae can store P in polyphosphate bodies through luxury-uptake; use
during deficiency
 HETEROTROPHY
• SOME ALGAE USE ORGANIC CARBON SOURCES IN THE DARK 
HETEROPHIC
• Hexoses are sterically incapable of diffusing across plasma membrane
passively
• Some algae posses an inducible active transport system through synthesis
of a glucose transport protein (30-4-kDa) which is induced in 15 min in
presence of glucose
• Process is energy dependent

• Amino-acids like arginine are actively transported across cell membranes

involving periplasmic binding proteins

• Small molecule-sized organic acids ( acetic acid, lactic acid) and ethanol,
can diffuse through cell membrane
• Free acids re more lipid soluble and less negatively charged than ionised
form; pH reduction favour their diffusion into cell

• Heterotrophic growth produces lower maximum specific growth rate than

photosynthetic growth  low affinity for the organic carbon
• Exception in Chlorella: heterotrophic and autotrophic growth rates are
comparable
 HETEROTROPHIC CULTURE
• growth on organic carbon sources : i) in the dark - heterotrophy
ii) in the light - mixotrophy

•Heterotrophy : advantages cf autotrophy:

i) can use high concentrations of org. C sources (without light) to produce high cell densities
ii) indoor cultures; operate in winter
ii) for producing high quality products

•Reactors of large volumes (500m3)

•Final conc. Of Chlorella can reach 100gL-1
•Doubling time reduced from 16 – 24h (phtoautotrophy) to 3.5 -4.8 h (heterotrophy)
•conditions
However application restricted to species that can grow and produce important metabolites under such

•Pigments not synthesised in heterotrophy

•PUFA not affected by heterotrophy
•Effects of mixotrophy under investigation; µmixotrophy = µautotrophy + µheterotrophy
•Heterotrophically grow Tetraselmis & Chlorella already in use as food for rotifers.
•investigated
Other species with higher DHA cont. like Nitzschia, Schizochytrium, Crypthecodinium being
 MIXOTROPHY
• Mixotrophy  Organic carbon and CO2 is simultaneously used in the light;
both respiratory & photosynthetic metabolism operating at the same time

• Mixotrophic growth rate = photoautotrophic growth rate + heterotrophic

growth rate

• Can achieve high cell concentration & productivity

• Increasing light intensity was found to increase utilisation of glucose in

Chlorella

• In mixotrophy, light energy is used for metabolism of the glucose or refixation

of CO2 into glucose through photosynthesis (see Richmond 2004; p. 120)