Phytoplankton

Phytoplankton (/ˌfaɪtoʊˈplæŋktən/) are the autotrophic (self-feeding) components of the plankton

community and a key part of ocean and freshwater ecosystems. The name comes from the Greek
words φυτόν (phyton), meaning 'plant', and πλαγκτός (planktos), meaning 'wanderer' or
'drifter'.[1][2][3]

Phytoplankton obtain their energy through photosynthesis, as trees and other plants do on land.
This means phytoplankton must have light from the sun, so they live in the well-lit surface layers
(euphotic zone) of oceans and lakes. In comparison with terrestrial plants, phytoplankton are
distributed over a larger surface area, are exposed to less seasonal variation and have markedly
faster turnover rates than trees (days versus decades). As a result, phytoplankton respond rapidly
on a global scale to climate variations.

Phytoplankton form the base of marine and freshwater food webs and are key players in the global
carbon cycle. They account for about half of global photosynthetic activity and at least half of the
oxygen production, despite amounting to only about 1% of the global plant biomass.

Phytoplankton are very diverse, comprising photosynthesizing bacteria (cyanobacteria) and various
unicellular protist groups (notably the diatoms).

Most phytoplankton are too small to be individually seen with the unaided eye. However, when
present in high enough numbers, some varieties may be noticeable as colored patches on the water
surface due to the presence of chlorophyll within their cells and accessory pigments (such as
phycobiliproteins or xanthophylls) in some species.

Types

Phytoplankton are photosynthesizing microscopic protists and bacteria that inhabit the upper sunlit
layer of marine and fresh water bodies of water on Earth. Paralleling plants on land, phytoplankton
undertake primary production in water,[2] creating organic compounds from carbon dioxide
dissolved in the water. Phytoplankton form the base of — and sustain — the aquatic food web,[4] and
are crucial players in the Earth's carbon cycle.[5]
 Diatoms are one of the most common types A cyanobacteria species (Cylindrospermum sp)
of phytoplankton

Phytoplankton are very diverse, comprising photosynthesizing bacteria (cyanobacteria) and various
unicellular protist groups (notably the diatoms). Many other organism groups formally named as
phytoplankton, including coccolithophores and dinoflagellates, are now no longer included as they
are not only phototrophic but can also eat.[6] These organisms are now more correctly
termed mixoplankton.[7] This recognition has important consequences for how we view the
functioning of the planktonic food web.[8]

Ecology

1:14

Global distribution of ocean phytoplankton – NASA

This visualization shows a model simulation of the dominant
phytoplankton types averaged over the period 1994–1998. * Red =
diatoms (big phytoplankton, which need silica) * Yellow = flagellates
(other big phytoplankton) * Green = prochlorococcus (small
phytoplankton that cannot use nitrate) * Cyan = synechococcus
(other small phytoplankton) Opacity indicates concentration of the
carbon biomass. In particular, the role of the swirls and filaments
(mesoscale features) appear important in maintaining high
biodiversity in the ocean.[5][9]

Phytoplankton obtain energy through the process of photosynthesis and must therefore live in the
well-lit surface layer (termed the euphotic zone) of an ocean, sea, lake, or other body of water.
Phytoplankton account for about half of all photosynthetic activity on Earth.[10][11][12] Their
cumulative energy fixation in carbon compounds (primary production) is the basis for the vast
majority of oceanic and also many freshwater food webs (chemosynthesis is a notable exception).

While almost all phytoplankton species are obligate photoautotrophs, there are some that are
mixotrophic and other, non-pigmented species that are actually heterotrophic (the latter are often
viewed as zooplankton).[2][13] Of these, the best known are dinoflagellate genera such as Noctiluca
and Dinophysis, that obtain organic carbon by ingesting other organisms or detrital material.

Phytoplankton live in the photic zone of the ocean, where photosynthesis is possible. During
photosynthesis, they assimilate carbon dioxide and release oxygen. If solar radiation is too high,
phytoplankton may fall victim to photodegradation. Phytoplankton species feature a large variety of
photosynthetic pigments which species-specifically enables them to absorb different wavelengths
of the variable underwater light.[14] This implies different species can use the wavelength of light
different efficiently and the light is not a single ecological resource but a multitude of resources
depending on its spectral composition.[15] By that it was found that changes in the spectrum of light
alone can alter natural phytoplankton communities even if the same intensity is available.[16] For
growth, phytoplankton cells additionally depend on nutrients, which enter the ocean by rivers,
continental weathering, and glacial ice meltwater on the poles. Phytoplankton release dissolved
organic carbon (DOC) into the ocean. Since phytoplankton are the basis of marine food webs, they
serve as prey for zooplankton, fish larvae and other heterotrophic organisms. They can also be
degraded by bacteria or by viral lysis. Although some phytoplankton cells, such as dinoflagellates,
are able to migrate vertically, they are still incapable of actively moving against currents, so they
slowly sink and ultimately fertilize the seafloor with dead cells and detritus.[17]
 Cycling of marine phytoplankton [17]

Phytoplankton are crucially dependent on a number of nutrients. These are primarily macronutrients
such as nitrate, phosphate or silicic acid, which are required in relatively large quantities for growth.
Their availability in the surface ocean is governed by the balance between the so-called biological
pump and upwelling of deep, nutrient-rich waters. The stoichiometric nutrient composition of
phytoplankton drives — and is driven by — the Redfield ratio of macronutrients generally available
throughout the surface oceans. However, across large areas of the oceans such as the Southern
Ocean, phytoplankton are limited by the lack of the micronutrient iron. This has led to some
scientists advocating iron fertilization as a means to counteract the accumulation of human-
produced carbon dioxide (CO2) in the atmosphere.[18] Large-scale experiments have added iron
(usually as salts such as ferrous sulfate) to the oceans to promote phytoplankton growth and draw
atmospheric CO2 into the ocean. Controversy about manipulating the ecosystem and the efficiency
of iron fertilization has slowed such experiments.[19]

Phytoplankton depend on B vitamins for survival. Areas in the ocean have been identified as having
a major lack of some B Vitamins, and correspondingly, phytoplankton.[20]

The effects of anthropogenic warming on the global population of phytoplankton is an area of active
research. Changes in the vertical stratification of the water column, the rate of temperature-
dependent biological reactions, and the atmospheric supply of nutrients are expected to have
important effects on future phytoplankton productivity.[21][22]
 Bioluminescence in phytoplankton
triggered by the agitation of waves
crashing on a beach

The effects of anthropogenic ocean acidification on phytoplankton growth and community structure
has also received considerable attention. The cells of coccolithophore phytoplankton are typically
covered in a calcium carbonate shell called a coccosphere that is sensitive to ocean acidification.
Because of their short generation times, evidence suggests some phytoplankton can adapt to
changes in pH induced by increased carbon dioxide on rapid time-scales (months to years).[23][24]

Phytoplankton serve as the base of the aquatic food web, providing an essential ecological function
for all aquatic life. Under future conditions of anthropogenic warming and ocean acidification,
changes in phytoplankton mortality due to changes in rates of zooplankton grazing may be
significant.[25] One of the many food chains in the ocean – remarkable due to the small number of
links – is that of phytoplankton sustaining krill (a crustacean similar to a tiny shrimp), which in turn
sustain baleen whales.

The El Niño-Southern Oscillation (ENSO) cycles in the Equatorial Pacific area can affect
phytoplankton.[26] Biochemical and physical changes during ENSO cycles modify the phytoplankton
community structure.[26] Also, changes in the structure of the phytoplankton, such as a significant
reduction in biomass and phytoplankton density, particularly during El Nino phases can occur.[27]
The sensitivity of phytoplankton to environmental changes is why they are often used as indicators
of estuarine and coastal ecological condition and health.[28] To study these events satellite ocean
color observations are used to observe these changes. Satellite images help to have a better view of
their global distribution.[26]

Diversity
 When two currents collide (here the
Oyashio and Kuroshio currents) they
create eddies. Phytoplankton
concentrates along the boundaries of
the eddies, tracing the motion of the
water.

Algal bloom off south west England

NASA satellite view of Southern

Ocean phytoplankton bloom

The term phytoplankton encompasses all photoautotrophic microorganisms in aquatic food webs.
However, unlike terrestrial communities, where most autotrophs are plants, phytoplankton are a
diverse group, incorporating protistan eukaryotes and both eubacterial and archaebacterial
prokaryotes. There are about 5,000 known species of marine phytoplankton.[29] How such diversity
evolved despite scarce resources (restricting niche differentiation) is unclear.[30]
In terms of numbers, the most important groups of phytoplankton include the diatoms,
cyanobacteria and dinoflagellates, although many other groups of algae are represented. One group,
the coccolithophorids, is responsible (in part) for the release of significant amounts of dimethyl
sulfide (DMS) into the atmosphere. DMS is oxidized to form sulfate which, in areas where ambient
aerosol particle concentrations are low, can contribute to the population of cloud condensation
nuclei, mostly leading to increased cloud cover and cloud albedo according to the so-called CLAW
hypothesis.[31][32] Different types of phytoplankton support different trophic levels within varying
ecosystems. In oligotrophic oceanic regions such as the Sargasso Sea or the South Pacific Gyre,
phytoplankton is dominated by the small sized cells, called picoplankton and nanoplankton (also
referred to as picoflagellates and nanoflagellates), mostly composed of cyanobacteria
(Prochlorococcus, Synechococcus) and picoeucaryotes such as Micromonas. Within more
productive ecosystems, dominated by upwelling or high terrestrial inputs, larger dinoflagellates are
the more dominant phytoplankton and reflect a larger portion of the biomass.[33]

Growth strategies

In the early twentieth century, Alfred C. Redfield found the similarity of the phytoplankton's elemental
composition to the major dissolved nutrients in the deep ocean.[34] Redfield proposed that the ratio
of carbon to nitrogen to phosphorus (106:16:1) in the ocean was controlled by the phytoplankton's
requirements, as phytoplankton subsequently release nitrogen and phosphorus as they are
remineralized. This so-called "Redfield ratio" in describing stoichiometry of phytoplankton and
seawater has become a fundamental principle to understand marine ecology, biogeochemistry and
phytoplankton evolution.[35] However, the Redfield ratio is not a universal value and it may diverge
due to the changes in exogenous nutrient delivery[36] and microbial metabolisms in the ocean, such
as nitrogen fixation, denitrification and anammox.

The dynamic stoichiometry shown in unicellular algae reflects their capability to store nutrients in an
internal pool, shift between enzymes with various nutrient requirements and alter osmolyte
composition.[37][38] Different cellular components have their own unique stoichiometry
characteristics,[35] for instance, resource (light or nutrients) acquisition machinery such as proteins
and chlorophyll contain a high concentration of nitrogen but low in phosphorus. Meanwhile, growth
machinery such as ribosomal RNA contains high nitrogen and phosphorus concentrations.

Based on allocation of resources, phytoplankton is classified into three different growth strategies,
namely survivalist, bloomer[39] and generalist. Survivalist phytoplankton has a high ratio of N:P (>30)
and contains an abundance of resource-acquisition machinery to sustain growth under scarce
resources. Bloomer phytoplankton has a low N:P ratio (<10), contains a high proportion of growth
machinery, and is adapted to exponential growth. Generalist phytoplankton has similar N:P to the
Redfield ratio and contain relatively equal resource-acquisition and growth machinery.

Factors affecting abundance

The NAAMES study was a five-year scientific research program conducted between 2015 and 2019
by scientists from Oregon State University and NASA to investigated aspects of phytoplankton
dynamics in ocean ecosystems, and how such dynamics influence atmospheric aerosols, clouds,
and climate (NAAMES stands for the North Atlantic Aerosols and Marine Ecosystems Study). The
study focused on the sub-arctic region of the North Atlantic Ocean, which is the site of one of
Earth's largest recurring phytoplankton blooms. The long history of research in this location, as well
as relative ease of accessibility, made the North Atlantic an ideal location to test prevailing scientific
hypotheses[40] in an effort to better understand the role of phytoplankton aerosol emissions on
Earth's energy budget.[41]

NAAMES was designed to target specific phases of the annual phytoplankton cycle: minimum,
climax and the intermediary decreasing and increasing biomass, in order to resolve debates on the
timing of bloom formations and the patterns driving annual bloom re-creation.[41] The NAAMES
project also investigated the quantity, size, and composition of aerosols generated by primary
production in order to understand how phytoplankton bloom cycles affect cloud formations and
climate.[42]
 Competing hypothesis of plankton variability[40]
Figure adapted from Behrenfeld & Boss 2014.[43]
Courtesy of NAAMES, Langley Research Center, NASA[44]

World concentrations of surface ocean chlorophyll as viewed by

satellite during the northern spring, averaged from 1998 to
2004. Chlorophyll is a marker for the distribution and abundance
of phytoplankton.
 Global patterns of monthly phytoplankton species richness and
species turnover
(A) Annual mean of monthly species richness and (B) month-to-
month species turnover projected by SDMs. Latitudinal
gradients of (C) richness and (D) turnover. Colored lines
(regressions with local polynomial fitting) indicate the means
per degree latitude from three different SDM algorithms used
(red shading denotes ±1 SD from 1000 Monte Carlo runs that
used varying predictors for GAM). Poleward of the thin
horizontal lines shown in (C) and (D), the model results cover
only <12 or <9 months, respectively.[45]

Factors affecting productivity

Environmental factors that affect phytoplankton productivity [46][47]

Phytoplankton are the key mediators of the biological pump. Understanding the response of
phytoplankton to changing environmental conditions is a prerequisite to predict future atmospheric
concentrations of CO2. Temperature, irradiance and nutrient concentrations, along with CO2 are the
chief environmental factors that influence the physiology and stoichiometry of phytoplankton.[48]
The stoichiometry or elemental composition of phytoplankton is of utmost importance to secondary
producers such as copepods, fish and shrimp, because it determines the nutritional quality and
influences energy flow through the marine food chains.[49] Climate change may greatly restructure
phytoplankton communities leading to cascading consequences for marine food webs, thereby
altering the amount of carbon transported to the ocean interior.[50][46]

The figure gives an overview of the various environmental factors that together affect phytoplankton
productivity. All of these factors are expected to undergo significant changes in the future ocean
due to global change.[51] Global warming simulations predict oceanic temperature increase;
dramatic changes in oceanic stratification, circulation and changes in cloud cover and sea ice,
resulting in an increased light supply to the ocean surface. Also, reduced nutrient supply is predicted
to co-occur with ocean acidification and warming, due to increased stratification of the water
column and reduced mixing of nutrients from the deep water to the surface.[52][46]

Role of phytoplankton

Role of phytoplankton on various compartments of the marine environment [53]

The compartments influenced by phytoplankton include the atmospheric gas composition,

inorganic nutrients, and trace element fluxes as well as the transfer and cycling of organic matter
via biological processes (see figure). The photosynthetically fixed carbon is rapidly recycled and
reused in the surface ocean, while a certain fraction of this biomass is exported as sinking particles
to the deep ocean, where it is subject to ongoing transformation processes, e.g., remineralization.[53]
Phytoplankton contribute to not only a basic pelagic marine food web but also to the microbial loop.
Phytoplankton are the base of the marine food web and because they do not rely on other
organisms for food, they make up the first trophic level. Organisms such as zooplankton feed on
these phytoplankton which are in turn fed on by other organisms and so forth until the fourth trophic
level is reached with apex predators. Approximately 90% of total carbon is lost between trophic
levels due to respiration, detritus, and dissolved organic matter. This makes the remineralization
process and nutrient cycling performed by phytoplankton and bacteria important in maintaining
efficiency.[54]

Phytoplankton blooms in which a species increases rapidly under conditions favorable to growth
can produce harmful algal blooms (HABs).

Aquaculture

Phytoplankton are a key food item in both aquaculture and mariculture. Both utilize phytoplankton
as food for the animals being farmed. In mariculture, the phytoplankton is naturally occurring and is
introduced into enclosures with the normal circulation of seawater. In aquaculture, phytoplankton
must be obtained and introduced directly. The plankton can either be collected from a body of water
or cultured, though the former method is seldom used. Phytoplankton is used as a foodstock for the
production of rotifers,[55] which are in turn used to feed other organisms. Phytoplankton is also used
to feed many varieties of aquacultured molluscs, including pearl oysters and giant clams. A 2018
study estimated the nutritional value of natural phytoplankton in terms of carbohydrate, protein and
lipid across the world ocean using ocean-colour data from satellites,[56] and found the calorific value
of phytoplankton to vary considerably across different oceanic regions and between different time
of the year.[56][57]

The production of phytoplankton under artificial conditions is itself a form of aquaculture.

Phytoplankton is cultured for a variety of purposes, including foodstock for other aquacultured
organisms,[55] a nutritional supplement for captive invertebrates in aquaria. Culture sizes range from
small-scale laboratory cultures of less than 1L to several tens of thousands of litres for commercial
aquaculture.[55] Regardless of the size of the culture, certain conditions must be provided for
efficient growth of plankton. The majority of cultured plankton is marine, and seawater of a specific
gravity of 1.010 to 1.026 may be used as a culture medium. This water must be sterilized, usually by
either high temperatures in an autoclave or by exposure to ultraviolet radiation, to prevent biological
contamination of the culture. Various fertilizers are added to the culture medium to facilitate the
growth of plankton. A culture must be aerated or agitated in some way to keep plankton suspended,
as well as to provide dissolved carbon dioxide for photosynthesis. In addition to constant aeration,
most cultures are manually mixed or stirred on a regular basis. Light must be provided for the
growth of phytoplankton. The colour temperature of illumination should be approximately 6,500 K,
but values from 4,000 K to upwards of 20,000 K have been used successfully. The duration of light
exposure should be approximately 16 hours daily; this is the most efficient artificial day length.[55]

Anthropogenic changes

Plot demonstrating increases in phytoplankton

species richness with increased temperature

Marine phytoplankton perform half of the global photosynthetic CO2 fixation (net global primary
production of ~50 Pg C per year) and half of the oxygen production despite amounting to only ~1%
of global plant biomass.[58] In comparison with terrestrial plants, marine phytoplankton are
distributed over a larger surface area, are exposed to less seasonal variation and have markedly
faster turnover rates than trees (days versus decades).[58] Therefore, phytoplankton respond rapidly
on a global scale to climate variations. These characteristics are important when one is evaluating
the contributions of phytoplankton to carbon fixation and forecasting how this production may
change in response to perturbations. Predicting the effects of climate change on primary
productivity is complicated by phytoplankton bloom cycles that are affected by both bottom-up
control (for example, availability of essential nutrients and vertical mixing) and top-down control (for
example, grazing and viruses).[59][58][60][61][62][63] Increases in solar radiation, temperature and
freshwater inputs to surface waters strengthen ocean stratification and consequently reduce
transport of nutrients from deep water to surface waters, which reduces primary
productivity.[58][63][64] Conversely, rising CO2 levels can increase phytoplankton primary production,
but only when nutrients are not limiting.[65][66][67][25]
Some studies indicate that overall global oceanic phytoplankton density has decreased in the past
century,[68] but these conclusions have been questioned because of the limited availability of long-
term phytoplankton data, methodological differences in data generation and the large annual and
decadal variability in phytoplankton production.[69][70][71][72] Moreover, other studies suggest a global
increase in oceanic phytoplankton production[73] and changes in specific regions or specific
phytoplankton groups.[74][75] The global Sea Ice Index is declining,[76] leading to higher light
penetration and potentially more primary production;[77] however, there are conflicting predictions
for the effects of variable mixing patterns and changes in nutrient supply and for productivity trends
in polar zones.[63][25]

The effect of human-caused climate change on phytoplankton biodiversity is not well understood.
Should greenhouse gas emissions continue rising to high levels by 2100, some phytoplankton
models predict an increase in species richness, or the number of different species within a given
area. This increase in plankton diversity is traced to warming ocean temperatures. In addition to
species richness changes, the locations where phytoplankton are distributed are expected to shift
towards the Earth's poles. Such movement may disrupt ecosystems, because phytoplankton are
consumed by zooplankton, which in turn sustain fisheries. This shift in phytoplankton location may
also diminish the ability of phytoplankton to store carbon that was emitted by human activities.
Human (anthropogenic) changes to phytoplankton impact both natural and economic processes.[78]

See also

Algaculture – Aquaculture involving the farming of algae

AlgaeBase – Species database

Algal bloom – Spread of planktonic algae in water

Bacterioplankton – Bacterial component of the plankton that drifts in the water column

Biological pump – Carbon capture process in oceans

CLAW hypothesis – A hypothesised negative feedback loop connecting the marine biota and the
climate

Critical depth

Deep chlorophyll maximum

Freshwater phytoplankton – Phytoplankton occurring in freshwater ecosystems

Iron fertilization – Ecological concept

 Microphyte – Microscopic algae (microalgae)

NAAMES

Ocean acidification – Decrease of pH levels in the ocean

Paradox of the plankton – The ecological observation of high plankton diversity despite
competition for few resources

Parasites of phytoplankton

Photosynthetic picoplankton – Group of photosynthetic plankton

Whiting event – Suspension of fine-grained calcium carbonate particles in water bodies

Thin layers (oceanography) – Congregations of plankton

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Further reading

Greeson, Phillip E. (1982). An annotated key to the identification of commonly occurring and
dominant genera of Algae observed in the Phytoplankton of the United States (https://pubs.er.usgs.
gov/publication/wsp2079) . Washington, D.C.: United States Government Printing Office.
ISBN 978-0-607-68844-3.

Kirby, Richard R. (2010). Ocean Drifters: A Secret World Beneath the Waves. Studio Cactus.
ISBN 978-1-904239-10-9.

Martin, Ronald; Quigg, Antonietta (2013). "Tiny Plants That Once Ruled the Seas". Scientific
American. 308 (6): 40–5. Bibcode:2013SciAm.308f..40M (https://ui.adsabs.harvard.edu/abs/201
3SciAm.308f..40M) . doi:10.1038/scientificamerican0613-40 (https://doi.org/10.1038%2Fscienti
ficamerican0613-40) . PMID 23729069 (https://pubmed.ncbi.nlm.nih.gov/23729069) .

External links

Secchi Disk and Secchi app (http://www.secchidisk.org) , a citizen science project to study the
phytoplankton

Ocean Drifters (https://vimeo.com/84872751/) , a short film narrated by David Attenborough

about the varied roles of plankton

Plankton Chronicles (http://www.planktonchronicles.org/en) , a short documentary films &

photos
DMS and Climate (https://web.archive.org/web/20150214081038/http://saga.pmel.noaa.gov/revi
ew/dms_climate.html) , NOAA

Plankton*Net (http://planktonnet.awi.de/) , images of planktonic species