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Biological oceanography: plankton and primary productivity

Marine Biology / Oceanography (B.Sc) · Ocean science · free study notes for exams, placements and interviews

In short

Biological oceanography studies the organisms of the open water column, how much organic carbon they create, and how that carbon moves through the food web and into the deep sea. Its core quantity is primary productivity — the rate at which phytoplankton convert dissolved CO₂ into organic matter by photosynthesis — because every fishery, every oxygen-minimum zone and the ocean's uptake of atmospheric CO₂ depends on it. In Indian waters the subject explains the monsoon-driven blooms of the Arabian Sea, the stratified low-productivity Bay of Bengal, the Kerala–Karnataka upwelling that supports the oil-sardine fishery, and recurring *Noctiluca* and *Trichodesmium* blooms. It is examined in B.Sc/M.Sc marine biology, oceanography and fisheries science papers, in CSIR-UGC NET (Earth, Atmospheric, Ocean and Planetary Sciences), ICAR-ARS and IIT JAM, and is standard interview ground for CSIR-NIO Goa, CMLRE and INCOIS (MoES), ICAR-CMFRI and CIFE, FSI, state fisheries departments and environmental consultancies doing marine EIA.

Explained

What plankton are, and how they are classified

The main groups

Primary productivity: definitions and controls

Measuring productivity and biomass

  1. Light-and-dark bottle (oxygen) method (Gaarder & Gran, 1927): paired clear and blackened BOD bottles incubated in situ at several depths; oxygen measured by Winkler titration or optode. Light bottle change = NPP, initial minus dark = respiration, light minus dark = GPP. Simple and calibration-free, but insensitive in oligotrophic water.
  2. ¹⁴C method (Steemann Nielsen, 1952): a known activity of NaH¹⁴CO₃ is added, incubated a few hours, filtered, and the particulate activity counted by liquid scintillation; dark-bottle counts are subtracted. Highly sensitive — the standard for open-ocean work, including on FORV *Sagar Sampada* — but needs a radiological licence; handling of radioisotopes in India is regulated by the atomic-energy regulator (the SHANTI Act, 2025 replaces the former Atomic Energy Act, 1962).
  3. ¹⁸O and ¹⁷O methods give true GPP; ¹⁵N uptake separates nitrate (new) from ammonium (regenerated) production and yields the f-ratio.
  4. Chlorophyll a: filter a known volume on GF/F, extract in 90% acetone, read on a spectrophotometer (trichromatic Jeffrey–Humphrey equations) or fluorometer (acidification method for phaeopigments); HPLC resolves marker pigments (fucoxanthin = diatoms, peridinin = dinoflagellates, zeaxanthin = cyanobacteria, 19'-hexanoyloxyfucoxanthin = prymnesiophytes). CHEMTAX apportions groups from pigments.
  5. In-situ and remote sensing: fluorometers on CTD rosettes and Argo/BGC-Argo floats, FRRF (variable fluorescence, Fv/Fm) for instantaneous photophysiology, flow cytometry for pico/nanoplankton, FlowCam and imaging systems for cell counts, eDNA metabarcoding for community composition. Ocean-colour satellites retrieve surface chlorophyll from water-leaving radiance ratios — India's OCM-2 (Oceansat-2) and OCM-3 (Oceansat-3/EOS-06, launched November 2022), processed by NRSC and used by INCOIS for the Potential Fishing Zone advisories; VGPM-type models convert satellite chlorophyll to depth-integrated production.
  6. Limitations: bottle incubations enclose and exclude large grazers ("bottle effect"), trace-metal contamination depresses rates, satellite chlorophyll sees only about the first optical depth and fails in turbid Case-2 coastal water, and chlorophyll-to-carbon ratios vary several-fold with light and nutrients.

Indian Ocean patterns, and the Indian institutional and legal frame

Key points

Formulas and examples

Light attenuation: I\_z = I₀ e−K\_d·z — I\_z = PAR at depth z (µmol photons m⁻² s⁻¹), K\_d = diffuse attenuation coefficient (m⁻¹), z in m.
Euphotic depth: Z\_eu = ln(100)/K\_d = 4.605/K\_d ; from a Secchi disc, K\_d ≈ 1.7/Z\_sd (Z\_sd = Secchi depth, m).
Oxygen method: NPP = (LB − IB)/t ; R = (IB − DB)/t ; GPP = (LB − DB)/t — LB, DB, IB = light, dark, initial O₂ (mg L⁻¹), t in hours.
O₂ to carbon: mg C = mg O₂ × (12/32) / PQ — PQ = photosynthetic quotient (mol O₂ evolved per mol CO₂ fixed), ≈1.0–1.2 (take 1.0 for nitrate-free assumptions unless stated).
¹⁴C method: PP (mg C m⁻³ per incubation) = (R\_s/R\_a) × ΣCO₂ ; rate = PP/t — R\_s = sample activity (DPM, dark-corrected), R\_a = activity added, ΣCO₂ = dissolved inorganic carbon (mg C m⁻³).
Chlorophyll a (Jeffrey & Humphrey trichromatic, 90% acetone, in µg mL⁻¹ of extract): C\_a = 11.85(E₆₆₄−E₇₅₀) − 1.54(E₆₄₇−E₇₅₀) − 0.08(E₆₃₀−E₇₅₀) ; then Chl a (mg m⁻³) = C\_a × v /(V × L) — v = extract volume (mL), V = volume filtered (L), L = cuvette path (cm).
Depth-integrated production: PP\_int = ∫₀Z\_eu PP(z) dz, evaluated by the trapezoidal rule over discrete depths (mg C m⁻² h⁻¹ → × day length for m⁻² d⁻¹).
Redfield stoichiometry: 106 CO₂ + 16 HNO₃ + H₃PO₄ + 122 H₂O → (CH₂O)₁₀₆(NH₃)₁₆H₃PO₄ + 138 O₂ ; C:N:P = 106:16:1 (atoms), C:N ≈ 6.6 by atoms, ≈ 5.7 by mass.
f-ratio: f = new production / (new + regenerated production), dimensionless, 0–1.
Nutrient uptake (Michaelis–Menten): µ = µ\_max·S/(K\_s + S) — S = nutrient concentration (µmol L⁻¹), K\_s = half-saturation constant.
VGPM (Behrenfeld & Falkowski): PP\_eu = 0.66125 × PB\_opt × [E₀/(E₀ + 4.1)] × Z\_eu × Chl\_sat × DL — PB\_opt in mg C (mg Chl)⁻¹ h⁻¹, E₀ = surface daily PAR (mol photons m⁻² d⁻¹), DL = day length (h).
Trophic transfer (Ryther): P\_fish = PP × En — E = transfer efficiency per step (≈0.1 in the open ocean, up to 0.15–0.2 in short upwelling chains), n = number of trophic steps.

Example 1 — light-and-dark bottle. Initial O₂ = 6.00 mg L⁻¹; after 4 h the light bottle reads 7.20 and the dark bottle 5.60 mg L⁻¹. Day length 12 h, PQ = 1.0.

Example 2 — ¹⁴C production and depth integration (upwelling station off Kochi). 1.0 × 10⁶ DPM of NaH¹⁴CO₃ added, dark-corrected filter activity 3,500 DPM, ΣCO₂ = 24,000 mg C m⁻³, incubation 4 h.

Example 3 — chlorophyll a. 2.0 L filtered, extracted in 10 mL of 90% acetone, 1-cm cuvette; E₆₆₄ = 0.045, E₆₄₇ = 0.012, E₆₃₀ = 0.008, E₇₅₀ = 0.002.

Example 4 — Secchi, euphotic depth and Redfield drawdown. Secchi depth 4.0 m.

Example 5 — satellite model and fish yield. Chl\_sat = 0.5 mg m⁻³, Z\_eu = 40 m, PB\_opt = 4.0, E₀ = 45 mol photons m⁻² d⁻¹, DL = 12 h.

FeatureDiatomsDinoflagellatesPicocyanobacteria
Size20–200 µm10–200 µm0.6–2 µm
MotilityNon-motileTwo flagellaNon-motile
WallSilica frustuleOrganic theca or nakedPeptidoglycan
Preferred waterCool, mixed, nutrient-richWarm, stratified, calmWarm, oligotrophic
Nutrient affinityLow (high K\_s), fast growthIntermediate, often mixotrophicVery high (low K\_s)
Fate of carbonSinks — high exportBlooms, often HABsMicrobial loop, low export
Indian example*Chaetoceros* in Kerala upwelling*Noctiluca* winter bloom, Arabian Sea*Prochlorococcus*, central Arabian Sea
Upwelling shelf (SW India, monsoon)Open Arabian Sea (inter-monsoon)Bay of Bengal (open)
Surface nitrateHigh (>5 µmol L⁻¹)Near zeroNear zero
Chlorophyll2–10 mg m⁻³0.1–0.3 mg m⁻³0.1–0.3 mg m⁻³
Production1–3 g C m⁻² d⁻¹0.2–0.5 g C m⁻² d⁻¹0.2–0.5 g C m⁻² d⁻¹
Dominant size classMicroplankton (diatoms)Pico/nanoplanktonPico/nanoplankton
f-ratioHigh (0.5–0.8)Low (0.1–0.2)Low
Limiting controlLight/stabilityNitrogenStratification (barrier layer)

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