Marine primary production

Source: Wikipedia, the free encyclopedia.

Ocean chlorophyll concentration as a proxy for marine primary production. Green indicates where there are a lot of phytoplankton, while blue indicates where there are few phytoplankton. – NASA Earth Observatory 2019.[1]

Marine primary production is the chemical synthesis in the ocean of

primary producers or autotrophs
.

Most marine primary production is generated by a diverse collection of

ocean food chain and produce half of the world's oxygen. Marine primary producers underpin almost all marine animal life by generating nearly all of the oxygen and food marine animals need to exist. Some marine primary producers are also ecosystem engineers which change the environment and provide habitats
for other marine life.

Primary production in the ocean can be contrasted with primary production on land. Globally the ocean and the land each produce about the same amount of primary production, but in the ocean primary production comes mainly from cyanobacteria and algae, while on land it comes mainly from vascular plants.

Marine algae includes the largely invisible and often

estuaries
. In addition, some seagrasses, like seaweeds, can be found at depths up to 50 metres on both soft and hard bottoms of the continental shelf.

Part of a series of overviews on
Marine life

Marine primary producers

Seasonal changes in which phytoplankton type dominates – NASA
click to animate
This visualization indicates seasonal changes in which phytoplankton types dominated 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.[2][3]
Part of a series on the
Carbon cycle
By regions
Carbon dioxide
Forms of carbon
Metabolic pathways
Carbon respiration
Carbon pumps
Carbon sequestration
Methane
Biogeochemical
Other

cold seeps and using chemosynthesis. However, most marine primary production comes from organisms which use photosynthesis on the carbon dioxide dissolved in the water. This process uses energy from sunlight to convert water and carbon dioxide[4]: 186–187  into sugars that can be used both as a source of chemical energy and of organic molecules that are used in the structural components of cells.[4]: 1242  Marine primary producers are important because they underpin almost all marine animal life by generating most of the oxygen
and food that provide other organisms with the chemical energy they need to exist.

The principal marine primary producers are cyanobacteria, algae and marine plants. The oxygen released as a by-product of photosynthesis is needed by nearly all living things to carry out cellular respiration. In addition, primary producers are influential in the global carbon and water cycles. They stabilize coastal areas and can provide habitats for marine animals. The term division has been traditionally used instead of phylum when discussing primary producers, although the International Code of Nomenclature for algae, fungi, and plants now accepts the terms as equivalent.[5]

In a reversal of the pattern on land, in the oceans, almost all photosynthesis is performed by algae and

red algae, and a diverse group of unicellular groups. Vascular plants are also represented in the ocean by groups such as the seagrasses
.

Unlike terrestrial ecosystems, the majority of primary production in the ocean is performed by free-living

littoral zone and adjacent shallow waters, where they can attach to the underlying substrate but still be within the photic zone. There are exceptions, such as Sargassum
, but the vast majority of free-floating production takes place within microscopic organisms.

The factors limiting primary production in the ocean are also very different from those on land. The availability of water, obviously, is not an issue (though its

nutrients, the building blocks for new growth, play crucial roles in regulating primary production in the ocean.[9] Available Earth System Models suggest that ongoing ocean bio-geochemical changes could trigger reductions in ocean NPP between 3% and 10% of current values depending on the emissions scenario.[10]

In 2020 researchers reported that measurements over the last two decades of primary production in the

carbon fixation in the future.[11][12]

Cyanobacteria

Evolution of photosynthesis from cyanobacteria
The origin and evolutionary tree of life that is based on small-subunit RNA. The branches that perform oxygenic photosynthesis are labeled with 'O2'. The black arrow indicates the plastid endosymbiotic event that resulted in the origin of eukaryotic photosynthesis from cyanobacteria-like organisms, which ultimately became chloroplasts in algae and later in plants. However, while chloroplasts of the higher plants, glaucophytes, green and red algae are thought to be the result of the plastid (primary) endosymbiosis, all other groups of algae are assumed to have arisen due to the algal (secondary and tertiary) endosymbiosis (not shown), in which one eukaryotic alga was incorporated into another eukaryote.[13][14][15][16][17] Only some branches of bacteria, eukarya, and archaea are displayed.[18]
Main article: Cyanobacteria

fix inorganic carbon into organic carbon compounds. They are found almost everywhere on earth: in damp soil, in both freshwater and marine environments, and even on Antarctic rocks.[19] In particular, some species occur as drifting cells floating in the ocean, and as such were amongst the first of the phytoplankton
. These bacteria function like algae in that they can process nitrogen from the atmosphere when none is in the ocean.

Cyanobacteria from a microbial mat. Cyanobacteria were the first organisms to release oxygen via photosynthesis.

The first primary producers that used photosynthesis were oceanic

dramatic change which redirected the evolution of the major animal and plant species.[22]

Prochlorococcus marinus

The tiny marine cyanobacterium Prochlorococcus, discovered in 1986, forms today part of the base of the ocean food chain and accounts for more than half the photosynthesis of the open ocean[23] and an estimated 20% of the oxygen in the Earth's atmosphere.[24] It is possibly the most plentiful genus on Earth: a single millilitre of surface seawater may contain 100,000 cells or more.[25]

Originally, biologists thought

prokaryotes, and hence cyanobacteria from the definition of algae.[26][27]

Helical cyanobacteria

Biological pigments

heterokonts
contain chlorophyll c instead of b, while red algae possess only chlorophyll a. All chlorophylls serve as the primary means plants use to intercept light in order to fuel photosynthesis.

Chloroplasts

Granum

(stack of
thylakoids)
Thylakoid lumen
Stroma
(aqueous fluid)
Parts of a chloroplast
Diagram above contains clickable links
Diagram above contains clickable links
Endosymbiosis
The first eukaryote may have originated from an ancestral prokaryote that had undergone membrane proliferation, compartmentalization of cellular function (into a nucleus, lysosomes, and an endoplasmic reticulum), and the establishment of endosymbiotic relationships with an aerobic prokaryote and, in some cases, a photosynthetic prokaryote to form mitochondria and chloroplasts, respectively.[30]

Chloroplasts (from the Greek chloros for green, and plastes for "the one who forms"[31]) are organelles that conduct photosynthesis, where the photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in the energy-storage molecules while freeing oxygen from water in plant and algal cells. They then use the stored energy to make organic molecules from carbon dioxide in a process known as the Calvin cycle.

A chloroplast is a type of organelle known as a

eukaryotic cell.[32]
Chloroplasts cannot be made by the plant cell and must be inherited by each daughter cell during cell division.

Most chloroplasts can probably be traced back to a single

tertiary endosymbiotic events
.

endosymbiotic origin from cyanobacteria.[33]

Microbial rhodopsin

Model of the energy generating mechanism in marine bacteria
      (1) When sunlight strikes a rhodopsin molecule
      (2) it changes its configuration so a proton is expelled from the cell
      (3) the chemical potential causes the proton to flow back to the cell
      (4) thus generating energy
      (5) in the form of adenosine triphosphate.[34]
Main article: Microbial rhodopsin

Phototrophic metabolism relies on one of three energy-converting pigments: chlorophyll, bacteriochlorophyll, and retinal. Retinal is the chromophore found in rhodopsins. The significance of chlorophyll in converting light energy has been written about for decades, but phototrophy based on retinal pigments is just beginning to be studied.[35]

salt evaporation ponds coloured purple by bacteriorhodopsin[36]
External videos
video icon Purple sulphur bacteria

In 2000 a team of microbiologists led by Edward DeLong made a crucial discovery in the understanding of the marine carbon and energy cycles. They discovered a gene in several species of bacteria[37][38] responsible for production of the protein rhodopsin, previously unheard of in bacteria. These proteins found in the cell membranes are capable of converting light energy to biochemical energy due to a change in configuration of the rhodopsin molecule as sunlight strikes it, causing the pumping of a proton from inside out and a subsequent inflow that generates the energy.[39] The archaeal-like rhodopsins have subsequently been found among different taxa, protists as well as in bacteria and archaea, though they are rare in complex multicellular organisms.[40][41][42]

Research in 2019 shows these "sun-snatching bacteria" are more widespread than previously thought and could change how oceans are affected by global warming. "The findings break from the traditional interpretation of marine ecology found in textbooks, which states that nearly all sunlight in the ocean is captured by chlorophyll in algae. Instead, rhodopsin-equipped bacteria function like hybrid cars, powered by organic matter when available — as most bacteria are — and by sunlight when nutrients are scarce."[43][35]

There is an astrobiological conjecture called the Purple Earth hypothesis which surmises that original life forms on Earth were retinal-based rather than chlorophyll-based, which would have made the Earth appear purple instead of green.[44][45]

Marine algae

Broad classification of algae [46]
Part of a series on
Plankton
Phytoplankton
Trophic mode
By size
By taxonomy
By habitat
  • Freshwater plankton
Other types
Blooms
Related topics

polyphyletic
. Unlike higher plants, algae lack roots, stems, or leaves.

Algal groups

Marine algae have traditionally been placed in groups such as: green algae, red algae, brown algae, diatoms, coccolithophores and dinoflagellates.

Green algae

Green algae live most of their lives as single cells or are filamentous, while others form colonies made up from long chains of cells, or are highly differentiated macroscopic seaweeds. They form an informal group containing about 8,000 recognized species.[47]

Red algae

Modern

multicellular with differentiated cells and include many notable seaweeds.[48][49] As coralline algae, they play an important role in the ecology of coral reefs. They form a (disputed) phylum containing about 7,000 recognized species.[48]

Brown algae

multicellular and include many seaweeds, including kelp. They form a class containing about 2,000 recognized species.[50]

Diatoms

Star stick diatom

Altogether, about 45 percent of the

diatoms.[51]

  • Diatoms are one of the most common types of phytoplankton
    Diatoms are one of the most common types of phytoplankton
  • They are a major algae group generating about 20% of world oxygen production.[52]
    They are a major algae group generating about 20% of world oxygen production.[52]
  • Diatoms have glass like cell walls called frustules which are made of silica.[53]
    Diatoms have glass like cell walls called
    silica.[53]
  • Diatoms linked in a colonial chain [54]
    Diatoms linked in a colonial chain [54]

Coccolithophores

marine food webs.[56] Management strategies are being employed to prevent eutrophication-related coccolithophore blooms, as these blooms lead to a decrease in nutrient flow to lower levels of the ocean.[57]

Dinoflagellate

Mixotrophic algae

Other groups

  • Diplonemids may be abundant in the world oceans
    Diplonemids
    may be abundant in the world oceans

Traditionally the

single cell genomics are being used in combination with high throughput techniques
.

Between 2009 and 2013, the

diplonemids. These organisms are generally colourless and oblong in shape, typically about 20 µm long and with two flagella.[61] Evidence from DNA barcoding suggests diplonemids may be among the most abundant and most species-rich of all marine eukaryote groups.[62][63]

By size

Algae can be classified by size as microalgae or macroalgae.

Microalgae

multicellular. Microalgae are important components of the marine protists, as well as the marine phytoplankton. They are very diverse. It has been estimated there are 200,000-800,000 species of which about 50,000 species have been described.[64]
Depending on the species, their sizes range from a few micrometers (µm) to a few hundred micrometers. They are specially adapted to an environment dominated by viscous forces.

Macroalgae

Kelp forests are among the most productive ecosystems on Earth.

multicellular and more visible types of algae, commonly called seaweeds. Seaweeds usually grow in shallow coastal waters where they are anchored to the seafloor by a holdfast. Seaweed that becomes adrift can wash up on beaches. Kelp is a large brown seaweed that forms large underwater forests covering about 25% of the world coastlines.[65] They are among the most productive and dynamic ecosystems on Earth.[66] Some Sargassum seaweeds are planktonic (free-floating) and form floating drifts.[67]: 246–255  Like microalgae, macroalgae (seaweeds) are technically marine protists
since they are not true plants.

brown or green algae
Global distribution of kelp forests
  • Macroalgae
  • Giant kelp is technically a protist since it is not a true plant, yet it is multicellular and can grow to 50 m
    Giant kelp
    is technically a protist since it is not a true plant, yet it is multicellular and can grow to 50 m
  • Sargassum seaweed is a brown alga with air bladders that help it float
    Sargassum seaweed is a brown alga with air bladders that help it float
  • Sargassum fish are camouflaged to live among drifting Sargassum seaweed
    Sargassum fish are camouflaged to live among drifting Sargassum seaweed
  • This unicellular bubble algae lives in tidal zones. It can have a 4 cm diameter.[68]
    This unicellular
    tidal zones. It can have a 4 cm diameter.[68]

Evolution of land plants

streptophytes [69]
Dating is roughly based on Morris et al. 2018.[70]

The diagram on the right shows an evolutionary scenario for the conquest of land by streptophytes.

parenchymatous — organism that formed rhizoidal structures and experienced desiccation from time to time. From this "hypothetical hydro-terrestrial alga", the lineages of Zygnematophyceae and embryophytes (land plants) arose.[69] In its infancy, the trajectory leading to the embryophytes was represented by the — now extinct — earliest land plants.[71]

The earliest land plants probably interacted with beneficial

tracheophytes evolved. While the exact trait repertoire of the hypothetical last common ancestor of land plants is uncertain, it will certainly have entailed properties of vascular and non-vascular plants. What is also certain is that the last common ancestor of land plants had traits of algal ancestry.[69]

Marine plants

Evolution of mangroves and seagrasses

Back in the

land plants we know today. Later, in the Cretaceous, some of these land plants returned to the sea as mangroves and seagrasses.[72]

Plant life can flourish in the brackish waters of

turtle grass
, Thalassia. These plants have adapted to the high salinity of the ocean environment.

Light is only able to penetrate the top 200 metres (660 ft) so this is the only part of the sea where plants can grow.[77] The surface layers are often deficient in biologically active nitrogen compounds. The marine nitrogen cycle consists of complex microbial transformations which include the fixation of nitrogen, its assimilation, nitrification, anammox and denitrification.[78] Some of these processes take place in deep water so that where there is an upwelling of cold waters, and also near estuaries where land-sourced nutrients are present, plant growth is higher. This means that the most productive areas, rich in plankton and therefore also in fish, are mainly coastal.[79]: 160–163 

Mangroves

Mangroves provide important nursery habitats for marine life, acting as hiding and foraging places for larval and juvenile forms of larger fish and invertebrates. Based on satellite data, the total world area of mangrove forests was estimated in 2010 as 134,257 square kilometres (51,837 sq mi).[80][81]

Global mangrove forests in 2000
common eelgrass according to IUCN data
  • Spalding, M. (2010) World atlas of mangroves, Routledge.
    doi:10.4324/9781849776608
    .

Seagrasses

Like mangroves, seagrasses provide important nursery habitats for larval and juvenile forms of larger fish and invertebrates. The total world area of seagrass meadows is more difficult to determine than mangrove forests, but was conservatively estimated in 2003 as 177,000 square kilometres (68,000 sq mi).[82]

  • Seagrass meadow
    Seagrass meadow
  • Sea dragons camouflaged to look like floating seaweed live in kelp forests and seagrass meadows[83]
    Sea dragons camouflaged to look like floating seaweed live in kelp forests and seagrass meadows[83]
External videos
video icon How did multicellularity evolve? – Journey to the Microcosmos

Stoichiometry

The

heterotrophic bacteria. In the face of global change, understanding and quantifying the mechanisms that lead to variability in C:N:P ratios are crucial in order to have an accurate projection of future climate change.[84]

Likely response of P:C and N:C ratios in marine phytoplankton
to major environmental drivers
Illustration of how the five environmental drivers under a typical future climate scenario affect the cellular allocation of volume between P-rich (red), N-rich (blue), and C-rich (orange) pools.[84]

A key unresolved question is what determines C:N:P of individual phytoplankton. Phytoplankton grows in the

acclimation), adaptation, and life history,[98][99] stoichiometric responses of phytoplankton can be variable even amongst closely related species.[84]

Meta-analysis/systematic review is a powerful statistical framework for synthesizing and integrating research results obtained from independent studies and for uncovering general trends.[100] The seminal synthesis by Geider and La Roche in 2002,[101] as well as the more recent work by Persson et al. in 2010,[102] has shown that C:P and N:P could vary by up to a factor of 20 between nutrient-replete and nutrient-limited cells. These studies have also shown that the C:N ratio can be modestly plastic due to nutrient limitation. A meta-analysis study by Hillebrand et al. in 2013 highlighted the importance of growth rate in determining elemental stoichiometry and showed that both C:P and N:P ratios decrease with the increasing growth rate.[103] In 2015, Yvon-Durocher et al. investigated the role of temperature in modulating C:N:P.[104] Although their dataset was limited to studies conducted prior to 1996, they have shown a statistically significant relationship between C:P and temperature increase. MacIntyre et al. (2002)[105] and Thrane et al. (2016)[106] have shown that irradiance plays an important role in controlling optimal cellular C:N and N:P ratios. Most recently, Moreno and Martiny (2018) provided a comprehensive summary of how environmental conditions regulate cellular stoichiometry from a physiological perspective.[93][84]

The elemental stoichiometry of marine phytoplankton plays a critical role in global biogeochemical cycles through its impact on nutrient cycling, secondary production, and carbon export. Although extensive laboratory experiments have been carried out over the years to assess the influence of different environmental drivers on the elemental composition of phytoplankton, a comprehensive quantitative assessment of the processes is still lacking. Here, the responses of P:C and N:C ratios of marine phytoplankton have been synthesized to five major drivers (inorganic phosphorus, inorganic nitrogen, inorganic iron, irradiance, and temperature) by a meta-analysis of experimental data across 366 experiments from 104 journal articles. These results show that the response of these ratios to changes in macronutrients is consistent across all the studies, where the increase in nutrient availability is positively related to changes in P:C and N:C ratios. The results show that eukaryotic phytoplankton are more sensitive to the changes in macronutrients compared to prokaryotes, possibly due to their larger cell size and their abilities to regulate their gene expression patterns quickly. The effect of irradiance was significant and constant across all studies, where an increase in irradiance decreased both P:C and N:C. The P:C ratio decreased significantly with warming, but the response to temperature changes was mixed depending on the culture growth mode and the growth phase at the time of harvest. Along with other oceanographic conditions of the subtropical gyres (e.g., low macronutrient availability), the elevated temperature may explain why P:C is consistently low in subtropical oceans. Iron addition did not systematically change either P:C or N:C.[84]

Evolutionary timeline

Evolutionary timeline of microbe-plant interactions
Microbial interactions outlined at the evolutionary scale, showing plant-microbe interactions occurring relatively recently compared to the more ancestral interactions among bacteria or between different microbial kingdoms. Both competitive (red) and cooperative (green) interactions within and between microbial kingdoms are depicted. Mya, million years ago.[107]
Evolutionary divergence estimated from Lücking et al., 2009 and Heckmanet al., 2001.[108][109]
Earth's biogeologic clock

See also

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Further reading