Marine primary production

Marine primary production is the chemical synthesis in the ocean of organic compounds from atmospheric or dissolved carbon dioxide. It principally occurs through the process of photosynthesis, which uses light as its source of energy, but it also occurs through chemosynthesis, which uses the oxidation or reduction of inorganic chemical compounds as its source of energy. Almost all life on Earth relies directly or indirectly on primary production. The organisms responsible for primary production are called primary producers or autotrophs.

Most marine primary production is generated by a diverse collection of marine microorganisms called algae and cyanobacteria. Together these form the principal primary producers at the base of the 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 unicellular microalgae, which together with cyanobacteria form the ocean phytoplankton, as well as the larger, more visible and complex multicellular macroalgae commonly called seaweed. Seaweeds are found along coastal areas, living on the floor of continental shelves and washed up in intertidal zones. Some seaweeds drift with plankton in the sunlit surface waters ( epipelagic zone) of the open ocean. Back in the Silurian, some phytoplankton evolved into red, brown and green algae. These algae then invaded the land and started evolving into the land plants we know today. Later in the Cretaceous some of these land plants returned to the sea as mangroves and seagrasses. These are found along coasts in intertidal regions and in the brackish water of 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.

Marine primary producers

Primary producers are the autotroph organisms that make their own food instead of eating other organisms. This means primary producers become the starting point in the food chain for heterotroph organisms that do eat other organisms. Some marine primary producers are specialised bacteria and archaea which are chemotrophs, making their own food by gathering around hydrothermal vents and 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 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. 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.

In a reversal of the pattern on land, in the oceans, almost all photosynthesis is performed by algae and cyanobacteria, with a small fraction contributed by vascular plants and other groups. Algae encompass a diverse range of organisms, ranging from single floating cells to attached seaweeds. They include photoautotrophs from a variety of groups. Eubacteria are important photosynthetizers in both oceanic and terrestrial ecosystems, and while some archaea are phototrophic, none are known to utilise oxygen-evolving photosynthesis. A number of eukaryotes are significant contributors to primary production in the ocean, including green algae, brown 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 microscopic organisms called phytoplankton. It has been estimated that half of the world's oxygen is produced by phytoplankton. Larger autotrophs, such as the seagrasses and macroalgae (seaweeds) are generally confined to the 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 salinity can be). Similarly, temperature, while affecting metabolic rates (see Q₁₀), ranges less widely in the ocean than on land because the heat capacity of seawater buffers temperature changes, and the formation of sea ice insulates it at lower temperatures. However, the availability of light, the source of energy for photosynthesis, and mineral nutrients, the building blocks for new growth, play crucial roles in regulating primary production in the ocean. 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.

In 2020 researchers reported that measurements over the last two decades of primary production in the Arctic Ocean show an increase of nearly 60% due to higher concentrations of phytoplankton. They hypothesize new nutrients are flowing in from other oceans and suggest this means the Arctic ocean may be able to support higher trophic level production and additional carbon fixation in the future.

Cyanobacteria

Cyanobacteria are a phylum (division) of bacteria, ranging from unicellular to filamentous and including colonial species, which 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. 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.

The first primary producers that used photosynthesis were oceanic cyanobacteria about 2.3 billion years ago. The release of molecular oxygen by cyanobacteria as a by-product of photosynthesis induced global changes in the Earth's environment. Because oxygen was toxic to most life on Earth at the time, this led to the near-extinction of oxygen-intolerant organisms, a dramatic change which redirected the evolution of the major animal and plant species.

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 and an estimated 20% of the oxygen in the Earth's atmosphere. It is possibly the most plentiful genus on Earth: a single millilitre of surface seawater may contain 100,000 cells or more.

Originally, biologists thought cyanobacteria was algae, and referred to it as "blue-green algae". The more recent view is that cyanobacteria are bacteria, and hence are not even in the same Kingdom as algae. Most authorities exclude all prokaryotes, and hence cyanobacteria from the definition of algae.

Biological pigments

Biological pigments are any coloured material in plant or animal cells. All biological pigments selectively absorb certain wavelengths of light while reflecting others.Lee, DW (2007) Nature's palette - the science of plant color. University of Chicago Press The primary function of pigments in plants is photosynthesis, which uses the green pigment chlorophyll and several colorful pigments that absorb as much light energy as possible. Chlorophyll is the primary pigment in plants; it is a chlorin that absorbs yellow and blue wavelengths of light while reflecting green. It is the presence and relative abundance of chlorophyll that gives plants their green color. Green algae and plants possess two forms of this pigment: chlorophyll ''a'' and chlorophyll ''b''. Kelps, diatoms, and other photosynthetic 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

Chloroplasts (from the Greek ''chloros'' for green, and ''plastes'' for "the one who forms") 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 plastid, characterized by its two membranes and a high concentration of chlorophyll. They are highly dynamic—they circulate and are moved around within plant cells, and occasionally pinch in two to reproduce. Their behavior is strongly influenced by environmental factors like light color and intensity. Chloroplasts, like mitochondria, contain their own DNA, which is thought to be inherited from their ancestor—a photosynthetic cyanobacterium that was engulfed by an early eukaryotic cell. 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 endosymbiotic event, when a cyanobacterium was engulfed by the eukaryote. Despite this, chloroplasts can be found in an extremely wide set of organisms, some not even directly related to each other—a consequence of many secondary and even tertiary endosymbiotic events.

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.

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 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.''Bacteria with Batteries'', Popular Science, January 2001, Page 55. 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.

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."

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.

Marine algae

Algae is an informal term for a widespread and diverse collection of photosynthetic eukaryotic organisms which are not necessarily closely related and are thus 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.

Red algae

Modern red algae are mostly multicellular with differentiated cells and include many notable seaweeds. 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.

File:Cyanidium O5A.jpg, Cyanidiophyceae colony, a class of unicellular red algae
File:Porphyra umbilicalis Helgoland.JPG, The seaweed ''Porphyra umbilicalis''

Brown algae

Brown algae are mostly multicellular and include many seaweeds, including kelp. They form a class containing about 2,000 recognized species.

Diatoms

Altogether, about 45 percent of the primary production in the oceans is contributed by diatoms.

File:Diatoms (248 05) Various diatoms.jpg, Diatoms are one of the most common types of phytoplankton
File:Diatoms through the microscope.jpg, They are a major algae group generating about 20% of world oxygen production.
File:Diatom algae Amphora sp.jpg, Diatoms have glass like cell walls called frustules which are made of silica.
File:Phytoplankton in the form of a diatom chain.jpg, Diatoms linked in a colonial chain

Coccolithophores

File:Emiliania huxleyi.png, The ubiquitous '' Emiliania huxleyi''
File:Cwall99 lg.jpg, ''Emiliania huxleyi'' bloom off south England

Coccolithophores are almost exclusively marine and are found in large numbers throughout the sunlight zone of the ocean. They have calcium carbonate plates (or scales) of uncertain function called ''coccoliths'', which are important microfossils. Coccolithophores are of interest to those studying global climate change because as ocean acidity increases, their coccoliths may become even more important as a carbon sink. The most abundant species of coccolithophore, '' Emiliania huxleyi'' is an ubiquitous component of the plankton base in marine food webs. 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.

Dinoflagellate

File:CSIRO ScienceImage 7609 SEM dinoflagellate.jpg, Dinoflagellates
File:Karenia brevis.jpg, ''Karenia brevis'' produces red tides highly toxic to humans
File:Algal bloom(akasio) by Noctiluca in Nagasaki.jpg, Red tide

Mixotrophic algae

Other groups

File:Diplonema papillatum SEM image.gif, Diplonemids may be abundant in the world oceans

Traditionally the phylogeny of microorganisms, such as the algal groups discussed above, was inferred and their taxonomy established based on studies of morphology. However developments in molecular phylogenetics have allowed the evolutionary relationship of species to be established by analyzing their DNA and protein sequences. Many taxa, including the algal groups discussed above, are in the process of being reclassified or redefined using molecular phylogenetics. Recent developments in molecular sequencing have allowed for the recovery of genomes directly from environmental samples and avoiding the need for culturing. This has led for example, to a rapid expansion in knowledge of the abundance and diversity of marine microorganisms. Molecular techniques such as genome-resolved metagenomics and single cell genomics are being used in combination with high throughput techniques.

Between 2009 and 2013, the ''Tara'' Oceans expedition traversed the world oceans collecting plankton and analysing them with contemporary molecular techniques. They found a huge range of previously unknown photosynthetic and mixotrophic algae. Among their findings were the diplonemids. These organisms are generally colourless and oblong in shape, typically about 20 µm long and with two flagella. Evidence from DNA barcoding suggests diplonemids may be among the most abundant and most species-rich of all marine eukaryote groups.

By size

Algae can be classified by size as ''microalgae'' or ''macroalgae''.

Microalgae

Microalgae are the microscopic types of algae, not visible to the naked eye. They are mostly unicellular species which exist as individuals or in chains or groups, though some are 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.Starckx, Senne (31 October 2012
A place in the sun - Algae is the crop of the future, according to researchers in Geel
Flanders Today, Retrieved 8 December 2012 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.

File:Zooxanthellae.jpg, Zooxanthellae is a photosynthetic algae that lives inside hosts like coral
File:Paramecium bursaria.jpg, A single-celled ciliate with green zoochlorellae living inside endosymbiotically
File:Euglena mutabilis - 400x - 1 (10388739803) (cropped).jpg, '' Euglena mutabilis'', a photosynthetic flagellate

Macroalgae

Macroalgae are the larger, 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. They are among the most productive and dynamic ecosystems on Earth.Mann, K.H. 1973. Seaweeds: their productivity and strategy for growth. Science 182: 975-981. Some ''Sargassum'' seaweeds are planktonic (free-floating) and form floating drifts. Like microalgae, macroalgae (seaweeds) are technically marine protists since they are not true plants.

File:Giant Kelp.jpg, Giant kelp is technically a protist since it is not a true plant, yet it is multicellular and can grow to 50 m
File:Sargassum on the beach, Cuba.JPG, ''Sargassum'' seaweed is a brown alga with air bladders that help it float

File:Histrio histrio by A. H. Baldwin.jpg, Sargassum fish are camouflaged to live among drifting ''Sargassum'' seaweed
File:Ventricaria ventricosa.JPG, This unicellular bubble algae lives in tidal zones. It can have a 4 cm diameter.

Evolution of land plants

The diagram on the right shows an evolutionary scenario for the conquest of land by streptophytes. Streptophyte algae include all green algae, and are the only photosynthetic eukaryotes from which the macroscopic land flora evolved ''(red lines)''. That said, throughout the course of evolution, algae from various other lineages have colonized land ''(yellow lines)'' — but also streptophyte algae have continuously and independently made the wet to dry transition (convergence of red and yellow). Throughout history, numerous lineages have become extinct ''(X labels)''. Terrestrial algae of various taxonomic affiliations dwell on rock surfaces and form biological soil crusts. From the diversity of the paraphyletic streptophyte algae, however, did an organism whose descendants eventually conquered land on a global scale emerge: a likely branched filamentous — or even 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. In its infancy, the trajectory leading to the embryophytes was represented by the — now extinct — earliest land plants.

The earliest land plants probably interacted with beneficial substrate microbiota that aided them in obtaining nutrients from their substrate. Furthermore, the earliest land plants had to successfully overcome a barrage of terrestrial stressors (including ultraviolet light and photosynthetically active irradiance, drought, drastic temperature shifts, etc.). They succeeded because they had the right set of traits — a mix of adaptations that were selected for in their hydro-terrestrial algal ancestors, exaptations, and the potential for co-option of a fortuitous set of genes and pathways. During the course of evolution, some members of the populations of the earliest land plants gained traits that are adaptive in terrestrial environments (such as some form of water conductance, stomata-like structures, embryos, etc.); eventually, the "hypothetical last common ancestor of land plants" emerged. From this ancestor, the extant bryophytes and 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.

Marine plants

Back in the Silurian, some phytoplankton evolved into red, brown and green algae. Green algae then invaded the land and started evolving into the land plants we know today. Later, in the Cretaceous, some of these land plants returned to the sea as mangroves and seagrasses.

Plant life can flourish in the brackish waters of estuaries, where mangroves or cordgrass or beach grass might grow. Flowering plants grow in sandy shallows in the form of seagrass meadows, mangroves line the coast in tropical and subtropical regions and salt-tolerant plants thrive in regularly inundated salt marshes. All of these habitats are able to sequester large quantities of carbon and support a biodiverse range of larger and smaller animal life. Marine plants can be found in intertidal zones and shallow waters, such as seagrasses like eelgrass

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