Primary production
Primary production is the production of organic compound
Organic compound
An organic compound is any member of a large class of gaseous, liquid, or solid chemical compounds whose molecules contain carbon. For historical reasons discussed below, a few types of carbon-containing compounds such as carbides, carbonates, simple oxides of carbon, and cyanides, as well as the...

s from atmospheric or aquatic carbon dioxide
Carbon dioxide
Carbon dioxide is a naturally occurring chemical compound composed of two oxygen atoms covalently bonded to a single carbon atom...

, principally through the process of photosynthesis
Photosynthesis is a chemical process that converts carbon dioxide into organic compounds, especially sugars, using the energy from sunlight. Photosynthesis occurs in plants, algae, and many species of bacteria, but not in archaea. Photosynthetic organisms are called photoautotrophs, since they can...

, with chemosynthesis
In biochemistry, chemosynthesis is the biological conversion of one or more carbon molecules and nutrients into organic matter using the oxidation of inorganic molecules or methane as a source of energy, rather than sunlight, as in photosynthesis...

 being much less important. Almost all life on earth is directly or indirectly reliant on primary production. The organisms responsible for primary production are known as primary producers or autotroph
An autotroph, or producer, is an organism that produces complex organic compounds from simple inorganic molecules using energy from light or inorganic chemical reactions . They are the producers in a food chain, such as plants on land or algae in water...

s, and form the base of the food chain
Food chain
A food web depicts feeding connections in an ecological community. Ecologists can broadly lump all life forms into one of two categories called trophic levels: 1) the autotrophs, and 2) the heterotrophs...

. In terrestrial ecoregions, these are mainly plant
Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...

s, while in aquatic ecoregions algae
Algae are a large and diverse group of simple, typically autotrophic organisms, ranging from unicellular to multicellular forms, such as the giant kelps that grow to 65 meters in length. They are photosynthetic like plants, and "simple" because their tissues are not organized into the many...

 are primarily responsible. Primary production is distinguished as either net or gross, the former accounting for losses to processes such as cellular respiration
Cellular respiration
Cellular respiration is the set of the metabolic reactions and processes that take place in the cells of organisms to convert biochemical energy from nutrients into adenosine triphosphate , and then release waste products. The reactions involved in respiration are catabolic reactions that involve...

, the latter not.


Primary production is the production of chemical energy
Chemical energy
Chemical energy is the potential of a chemical substance to undergo a transformation through a chemical reaction or, to transform other chemical substances...

 in organic compounds by living organism
In biology, an organism is any contiguous living system . In at least some form, all organisms are capable of response to stimuli, reproduction, growth and development, and maintenance of homoeostasis as a stable whole.An organism may either be unicellular or, as in the case of humans, comprise...

s. The main source of this energy is sunlight
Sunlight, in the broad sense, is the total frequency spectrum of electromagnetic radiation given off by the Sun. On Earth, sunlight is filtered through the Earth's atmosphere, and solar radiation is obvious as daylight when the Sun is above the horizon.When the direct solar radiation is not blocked...

 but a minute fraction of primary production is driven by lithotroph
A lithotroph is an organism that uses an inorganic substrate to obtain reducing equivalents for use in biosynthesis or energy conservation via aerobic or anaerobic respiration. Known chemolithotrophs are exclusively microbes; No known macrofauna possesses the ability to utilize inorganic...

ic organisms using the chemical energy of inorganic molecules.

Regardless of its source, this energy is used to synthesize complex organic molecules from simpler inorganic compounds such as carbon dioxide
Carbon dioxide
Carbon dioxide is a naturally occurring chemical compound composed of two oxygen atoms covalently bonded to a single carbon atom...

 (CO2) and water
Water is a chemical substance with the chemical formula H2O. A water molecule contains one oxygen and two hydrogen atoms connected by covalent bonds. Water is a liquid at ambient conditions, but it often co-exists on Earth with its solid state, ice, and gaseous state . Water also exists in a...

 (H2O). The following two equations are simplified representations of photosynthesis (top) and (one form of) chemosynthesis
In biochemistry, chemosynthesis is the biological conversion of one or more carbon molecules and nutrients into organic matter using the oxidation of inorganic molecules or methane as a source of energy, rather than sunlight, as in photosynthesis...

 (bottom) :
CO2 + H2O + light CH2O + O2
CO2 + O2 + 4 H2S CH2O + 4 S + 3 H2O

In both cases, the end point is reduced
Redox reactions describe all chemical reactions in which atoms have their oxidation state changed....

A carbohydrate is an organic compound with the empirical formula ; that is, consists only of carbon, hydrogen, and oxygen, with a hydrogen:oxygen atom ratio of 2:1 . However, there are exceptions to this. One common example would be deoxyribose, a component of DNA, which has the empirical...

 (CH2O), typically molecules such as glucose
Glucose is a simple sugar and an important carbohydrate in biology. Cells use it as the primary source of energy and a metabolic intermediate...

 or other sugar
Sugar is a class of edible crystalline carbohydrates, mainly sucrose, lactose, and fructose, characterized by a sweet flavor.Sucrose in its refined form primarily comes from sugar cane and sugar beet...

s. These relatively simple molecules may be then used to further synthesise more complicated molecules, including protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...

s, complex carbohydrates
Complex carbohydrates
Complex carbohydrates may refer to:*oligosaccharides, a saccharide polymer containing a small number of component sugars...

, lipids, and nucleic acids, or be respired
Cellular respiration
Cellular respiration is the set of the metabolic reactions and processes that take place in the cells of organisms to convert biochemical energy from nutrients into adenosine triphosphate , and then release waste products. The reactions involved in respiration are catabolic reactions that involve...

 to perform work
Work (thermodynamics)
In thermodynamics, work performed by a system is the energy transferred to another system that is measured by the external generalized mechanical constraints on the system. As such, thermodynamic work is a generalization of the concept of mechanical work in mechanics. Thermodynamic work encompasses...

. Consumption of primary producers by heterotroph
A heterotroph is an organism that cannot fix carbon and uses organic carbon for growth. This contrasts with autotrophs, such as plants and algae, which can use energy from sunlight or inorganic compounds to produce organic compounds such as carbohydrates, fats, and proteins from inorganic carbon...

ic organisms, such as animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...

s, then transfers these organic molecules (and the energy stored within them) up the food web
Food web
A food web depicts feeding connections in an ecological community. Ecologists can broadly lump all life forms into one of two categories called trophic levels: 1) the autotrophs, and 2) the heterotrophs...

, fueling all of the Earth
Earth is the third planet from the Sun, and the densest and fifth-largest of the eight planets in the Solar System. It is also the largest of the Solar System's four terrestrial planets...

's living systems.


Gross primary production (GPP) is the rate at which an ecosystem's producers capture and store a given amount of chemical energy as biomass in a given length of time. Some fraction of this fixed energy is used by primary producers for cellular respiration
Cellular respiration
Cellular respiration is the set of the metabolic reactions and processes that take place in the cells of organisms to convert biochemical energy from nutrients into adenosine triphosphate , and then release waste products. The reactions involved in respiration are catabolic reactions that involve...

 and maintenance of existing tissues (i.e., "growth respiration" and "maintenance respiration
Maintenance respiration
Maintenance respiration refers to metabolism occurring in an organism that is needed to maintain that organism in a healthy, living state...

"). The remaining fixed energy (i.e., mass of photosynthate) is referred to as net primary production (NPP).
NPP = GPP - respiration [by plants]

Net primary production is the rate at which all the plants in an ecosystem produce net useful chemical energy; it is equal to the difference between the rate at which the plants in an ecosystem produce useful chemical energy (GPP) and the rate at which they use some of that energy during respiration. Some net primary production goes toward growth and reproduction of primary producers, while some is consumed by herbivores.

Both gross and net primary production are in units of mass / area / time. In terrestrial ecosystems, mass of carbon per unit area per year (g C/m2/yr) is most often used as the unit of measurement.

Terrestrial production

On the land, almost all primary production is now performed by vascular plant
Vascular plant
Vascular plants are those plants that have lignified tissues for conducting water, minerals, and photosynthetic products through the plant. Vascular plants include the clubmosses, Equisetum, ferns, gymnosperms and angiosperms...

s, with a small fraction coming from algae and non-vascular plant
Non-vascular plant
Non-vascular plants is a general term for those plants without a vascular system . Although non-vascular plants lack these particular tissues, a number of non-vascular plants possess tissues specialized for internal transport of water....

s such as moss
Mosses are small, soft plants that are typically 1–10 cm tall, though some species are much larger. They commonly grow close together in clumps or mats in damp or shady locations. They do not have flowers or seeds, and their simple leaves cover the thin wiry stems...

es and liverworts
The Marchantiophyta are a division of bryophyte plants commonly referred to as hepatics or liverworts. Like other bryophytes, they have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of genetic information....

. Before the evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...

 of vascular plants, non-vascular plants likely played a more significant role. Primary production on land is a function
Function (mathematics)
In mathematics, a function associates one quantity, the argument of the function, also known as the input, with another quantity, the value of the function, also known as the output. A function assigns exactly one output to each input. The argument and the value may be real numbers, but they can...

 of many factors, but principally local hydrology
Hydrology is the study of the movement, distribution, and quality of water on Earth and other planets, including the hydrologic cycle, water resources and environmental watershed sustainability...

 and temperature
Temperature is a physical property of matter that quantitatively expresses the common notions of hot and cold. Objects of low temperature are cold, while various degrees of higher temperatures are referred to as warm or hot...

 (the latter covaries to an extent with light, the source of energy for photosynthesis). While plants cover much of the Earth's surface, they are strongly curtailed wherever temperatures are too extreme or where necessary plant resources (principally water and light) are limiting, such as desert
A desert is a landscape or region that receives an extremely low amount of precipitation, less than enough to support growth of most plants. Most deserts have an average annual precipitation of less than...

s or polar region
Polar region
Earth's polar regions are the areas of the globe surrounding the poles also known as frigid zones. The North Pole and South Pole being the centers, these regions are dominated by the polar ice caps, resting respectively on the Arctic Ocean and the continent of Antarctica...


Water is "consumed" in plants by the processes of photosynthesis (see above) and transpiration
Transpiration is a process similar to evaporation. It is a part of the water cycle, and it is the loss of water vapor from parts of plants , especially in leaves but also in stems, flowers and roots. Leaf surfaces are dotted with openings which are collectively called stomata, and in most plants...

. The latter process (which is responsible for about 90% of water use) is driven by the evaporation
Evaporation is a type of vaporization of a liquid that occurs only on the surface of a liquid. The other type of vaporization is boiling, which, instead, occurs on the entire mass of the liquid....

 of water from the leaves
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....

 of plants. Transpiration allows plants to transport water and mineral
A mineral is a naturally occurring solid chemical substance formed through biogeochemical processes, having characteristic chemical composition, highly ordered atomic structure, and specific physical properties. By comparison, a rock is an aggregate of minerals and/or mineraloids and does not...

A nutrient is a chemical that an organism needs to live and grow or a substance used in an organism's metabolism which must be taken in from its environment. They are used to build and repair tissues, regulate body processes and are converted to and used as energy...

s from the soil
Soil is a natural body consisting of layers of mineral constituents of variable thicknesses, which differ from the parent materials in their morphological, physical, chemical, and mineralogical characteristics...

 to growth regions, and also cools the plant. Diffusion of water out of a leaf, the force that drives transpiration, is regulated by structures known as stoma
In botany, a stoma is a pore, found in the leaf and stem epidermis that is used forgas exchange. The pore is bordered by a pair of specialized parenchyma cells known as guard cells that are responsible for regulating the size of the opening...

ta. These also regulate the diffusion of carbon dioxide from the atmosphere into the leaf, such that decreasing water loss (by partially closing stomata) also decreases carbon dioxide gain. Certain plants use alternative forms of photosynthesis, called Crassulacean acid metabolism
Crassulacean acid metabolism
Crassulacean acid metabolism, also known as CAM photosynthesis, is a carbon fixation pathway that evolved in some plants as an adaptation to arid conditions. The stomata in the leaves remain shut during the day to reduce evapotranspiration, but open at night to collect carbon dioxide...

 (CAM) and C4
C4 carbon fixation
C4 carbon fixation is one of three biochemical mechanisms, along with and CAM photosynthesis, used in carbon fixation. It is named for the 4-carbon molecule present in the first product of carbon fixation in these plants, in contrast to the 3-carbon molecule products in plants. fixation is an...

. These employ physiological
Physiology is the science of the function of living systems. This includes how organisms, organ systems, organs, cells, and bio-molecules carry out the chemical or physical functions that exist in a living system. The highest honor awarded in physiology is the Nobel Prize in Physiology or...

 and anatomical
Anatomy is a branch of biology and medicine that is the consideration of the structure of living things. It is a general term that includes human anatomy, animal anatomy , and plant anatomy...

 adaptations to increase water-use efficiency and allow increased primary production to take place under conditions that would normally limit carbon fixation by C3
C3 carbon fixation
carbon fixation is a metabolic pathway for carbon fixation in photosynthesis. This process converts carbon dioxide and ribulose bisphosphate into 3-phosphoglycerate through the following reaction:...

 plants (the majority of plant species).

Oceanic production

In a reversal of the pattern on land, in the oceans, almost all primary production is performed by algae, 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
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...

 are important photosynthetizers in both oceanic and terrestrial ecosystems, and while some archaea
The Archaea are a group of single-celled microorganisms. A single individual or species from this domain is called an archaeon...

 are phototroph
Phototrophs are the organisms that carry out photosynthesis to acquire energy. They use the energy from sunlight to convert carbon dioxide and water into organic material to be utilized in cellular functions such as biosynthesis and respiration.Most phototrophs are autotrophs, also known as...

ic, none are known to utilise oxygen-evolving photosynthesis. A number of eukaryote
A eukaryote is an organism whose cells contain complex structures enclosed within membranes. Eukaryotes may more formally be referred to as the taxon Eukarya or Eukaryota. The defining membrane-bound structure that sets eukaryotic cells apart from prokaryotic cells is the nucleus, or nuclear...

s 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 seagrass
Seagrasses are flowering plants from one of four plant families , all in the order Alismatales , which grow in marine, fully saline environments.-Ecology:...


Unlike terrestrial ecosystems, the majority of primary production in the ocean is performed by free-living microscopic organisms
A microorganism or microbe is a microscopic organism that comprises either a single cell , cell clusters, or no cell at all...

 called phytoplankton
Phytoplankton are the autotrophic component of the plankton community. The name comes from the Greek words φυτόν , meaning "plant", and πλαγκτός , meaning "wanderer" or "drifter". Most phytoplankton are too small to be individually seen with the unaided eye...

. Larger autotrophs, such as the seagrasses and macroalgae (seaweed
Seaweed is a loose, colloquial term encompassing macroscopic, multicellular, benthic marine algae. The term includes some members of the red, brown and green algae...

s) are generally confined to the littoral
The littoral zone is that part of a sea, lake or river that is close to the shore. In coastal environments the littoral zone extends from the high water mark, which is rarely inundated, to shoreline areas that are permanently submerged. It always includes this intertidal zone and is often used to...

 zone and adjacent shallow waters, where they can attach
A holdfast is a root-like structure that anchors aquatic sessile organisms, such as seaweed, other sessile algae, stalked crinoids, benthic cnidarians, and sponges, to the substrate. ...

 to the underlying substrate but still be within the photic zone
Photic zone
The photic zone or euphotic zone is the depth of the water in a lake or ocean that is exposed to sufficient sunlight for photosynthesis to occur...

. There are exceptions, such as Sargassum
Sargassum is a genus of brown macroalga in the order Fucales. Numerous species are distributed throughout the temperate and tropical oceans of the world, where they generally inhabit shallow water and coral reefs. However, the genus may be best known for its planktonic species...

, 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
Salinity is the saltiness or dissolved salt content of a body of water. It is a general term used to describe the levels of different salts such as sodium chloride, magnesium and calcium sulfates, and bicarbonates...

 can be). Similarly, temperature, while affecting metabolic
Metabolism is the set of chemical reactions that happen in the cells of living organisms to sustain life. These processes allow organisms to grow and reproduce, maintain their structures, and respond to their environments. Metabolism is usually divided into two categories...

 rates (see Q10
Q10 (temperature coefficient)
The Q10 temperature coefficient is a measure of the rate of change of a biological or chemical system as a consequence of increasing the temperature by 10 °C. There are many examples where the Q10 is used, one being the calculation of the nerve conduction velocity and another being calculating the...

), ranges less widely in the ocean than on land because the heat capacity
Heat capacity
Heat capacity , or thermal capacity, is the measurable physical quantity that characterizes the amount of heat required to change a substance's temperature by a given amount...

 of seawater buffers temperature changes, and the formation of sea ice
Sea ice
Sea ice is largely formed from seawater that freezes. Because the oceans consist of saltwater, this occurs below the freezing point of pure water, at about -1.8 °C ....

Thermal insulation
Thermal insulation is the reduction of the effects of the various processes of heat transfer between objects in thermal contact or in range of radiative influence. Heat transfer is the transfer of thermal energy between objects of differing temperature...

 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.


The sunlit zone of the ocean is called the photic zone
Photic zone
The photic zone or euphotic zone is the depth of the water in a lake or ocean that is exposed to sufficient sunlight for photosynthesis to occur...

 (or euphotic zone). This is a relatively thin layer (10–100 m) near the ocean's surface where there is sufficient light for photosynthesis to occur. For practical purposes, the thickness of the photic zone is typically defined by the depth at which light reaches 1% of its surface value. Light is attenuated
In physics, attenuation is the gradual loss in intensity of any kind of flux through a medium. For instance, sunlight is attenuated by dark glasses, X-rays are attenuated by lead, and light and sound are attenuated by water.In electrical engineering and telecommunications, attenuation affects the...

 down the water column by its absorption or scattering
Scattering is a general physical process where some forms of radiation, such as light, sound, or moving particles, are forced to deviate from a straight trajectory by one or more localized non-uniformities in the medium through which they pass. In conventional use, this also includes deviation of...

 by the water itself, and by dissolved or particulate material within it (including phytoplankton).

Net photosynthesis in the water column is determined by the interaction between the photic zone and the mixed layer
Mixed layer
The oceanic or limnological mixed layer is a layer in which active turbulence has homogenized some range of depths. The surface mixed layer is a layer where this turbulence is generated by winds, cooling, or processes such as evaporation or sea ice formation which result in an increase in salinity...

. Turbulent mixing
In fluid dynamics, turbulence or turbulent flow is a flow regime characterized by chaotic and stochastic property changes. This includes low momentum diffusion, high momentum convection, and rapid variation of pressure and velocity in space and time...

 by wind
Wind is the flow of gases on a large scale. On Earth, wind consists of the bulk movement of air. In outer space, solar wind is the movement of gases or charged particles from the sun through space, while planetary wind is the outgassing of light chemical elements from a planet's atmosphere into space...

 energy at the ocean's surface homogenises the water column vertically until the turbulence dissipates
In physics, dissipation embodies the concept of a dynamical system where important mechanical models, such as waves or oscillations, lose energy over time, typically from friction or turbulence. The lost energy converts into heat, which raises the temperature of the system. Such systems are called...

 (creating the aforementioned mixed layer). The deeper the mixed layer, the lower the average amount of light intercepted by phytoplankton within it. The mixed layer can vary from being shallower than the photic zone, to being much deeper than the photic zone. When it is much deeper than the photic zone, this results in phytoplankton spending too much time in the dark for net growth to occur. The maximum depth of the mixed layer in which net growth can occur is called the critical depth
Critical Depth
In biological oceanography, Critical Depth is defined as a hypothesized surface mixing depth at which phytoplankton growth is precisely matched by losses of phytoplankton biomass within this depth interval. -History:...

. As long as there are adequate nutrients available, net primary production occurs whenever the mixed layer is shallower than the critical depth.

Both the magnitude of wind mixing and the availability of light at the ocean's surface are affected across a range of space- and time-scales. The most characteristic of these is the seasonal cycle
A season is a division of the year, marked by changes in weather, ecology, and hours of daylight.Seasons result from the yearly revolution of the Earth around the Sun and the tilt of the Earth's axis relative to the plane of revolution...

 (caused by the consequences
Effect of sun angle on climate
The amount of heat energy received at any location on the globe is a direct effect of sun angle on climate, as the angle at which sunlight strikes the Earth varies by location, time of day, and season due to the Earth's orbit around the sun and the Earth's rotation around its tilted axis...

 of the Earth's axial tilt
Axial tilt
In astronomy, axial tilt is the angle between an object's rotational axis, and a line perpendicular to its orbital plane...

), although wind magnitudes additionally have strong spatial components. Consequently, primary production in temperate
In geography, temperate or tepid latitudes of the globe lie between the tropics and the polar circles. The changes in these regions between summer and winter are generally relatively moderate, rather than extreme hot or cold...

 regions such as the North Atlantic is highly seasonal, varying with both incident light at the water's surface (reduced in winter) and the degree of mixing (increased in winter). In tropical
The tropics is a region of the Earth surrounding the Equator. It is limited in latitude by the Tropic of Cancer in the northern hemisphere at approximately  N and the Tropic of Capricorn in the southern hemisphere at  S; these latitudes correspond to the axial tilt of the Earth...

 regions, such as the gyre
A gyre in oceanography is any large system of rotating ocean currents, particularly those involved with large wind movements. Gyres are caused by the Coriolis Effect; planetary vorticity along with horizontal and vertical friction, which determine the circulation patterns from the wind curl...

s in the middle of the major basins
Oceanic basin
Hydrologically, an oceanic basin may be anywhere on Earth that is covered by seawater, but geologically ocean basins are large geologic basins that are below sea level...

, light may only vary slightly across the year, and mixing may only occur episodically, such as during large storm
A storm is any disturbed state of an astronomical body's atmosphere, especially affecting its surface, and strongly implying severe weather...

s or hurricanes.


Mixing also plays an important role in the limitation of primary production by nutrients. Inorganic nutrients, such as nitrate
The nitrate ion is a polyatomic ion with the molecular formula NO and a molecular mass of 62.0049 g/mol. It is the conjugate base of nitric acid, consisting of one central nitrogen atom surrounded by three identically-bonded oxygen atoms in a trigonal planar arrangement. The nitrate ion carries a...

, phosphate
A phosphate, an inorganic chemical, is a salt of phosphoric acid. In organic chemistry, a phosphate, or organophosphate, is an ester of phosphoric acid. Organic phosphates are important in biochemistry and biogeochemistry or ecology. Inorganic phosphates are mined to obtain phosphorus for use in...

 and silicic acid
Silicic acid
Silicic acid is a general name for a family of chemical compounds of the element silicon, hydrogen, and oxygen, with the general formula [SiOx4-2x]n...

 are necessary for phytoplankton to synthesis
Organic synthesis
Organic synthesis is a special branch of chemical synthesis and is concerned with the construction of organic compounds via organic reactions. Organic molecules can often contain a higher level of complexity compared to purely inorganic compounds, so the synthesis of organic compounds has...

e their cells and cellular machinery. Because of gravitational sinking of particulate material (such as plankton
Plankton are any drifting organisms that inhabit the pelagic zone of oceans, seas, or bodies of fresh water. That is, plankton are defined by their ecological niche rather than phylogenetic or taxonomic classification...

, dead or fecal material), nutrients are constantly lost from the photic zone, and are only replenished by mixing or upwelling
Upwelling is an oceanographic phenomenon that involves wind-driven motion of dense, cooler, and usually nutrient-rich water towards the ocean surface, replacing the warmer, usually nutrient-depleted surface water. The increased availability in upwelling regions results in high levels of primary...

 of deeper water. This is exacerbated where summertime solar heating and reduced winds increases vertical stratification and leads to a strong thermocline
A thermocline is a thin but distinct layer in a large body of fluid , in which temperature changes more rapidly with depth than it does in the layers above or below...

, since this makes it more difficult for wind mixing to entrain deeper water. Consequently, between mixing events, primary production (and the resulting processes that leads to sinking particulate material) constantly acts to consume nutrients in the mixed layer, and in many regions this leads to nutrient exhaustion and decreased mixed layer production in the summer (even in the presence of abundant light). However, as long as the photic zone is deep enough, primary production may continue below the mixed layer where light-limited growth rates mean that nutrients are often more abundant.


Another factor relatively recently discovered to play a significant role in oceanic primary production is the micronutrient
Micronutrients are nutrients required by humans and other living things throughout life in small quantities to orchestrate a whole range of physiological functions, but which the organism itself cannot produce. For people, they include dietary trace minerals in amounts generally less than 100...

Iron is a chemical element with the symbol Fe and atomic number 26. It is a metal in the first transition series. It is the most common element forming the planet Earth as a whole, forming much of Earth's outer and inner core. It is the fourth most common element in the Earth's crust...

. This is used as a cofactor
Cofactor (biochemistry)
A cofactor is a non-protein chemical compound that is bound to a protein and is required for the protein's biological activity. These proteins are commonly enzymes, and cofactors can be considered "helper molecules" that assist in biochemical transformations....

 in enzyme
Enzymes are proteins that catalyze chemical reactions. In enzymatic reactions, the molecules at the beginning of the process, called substrates, are converted into different molecules, called products. Almost all chemical reactions in a biological cell need enzymes in order to occur at rates...

s involved in processes such as nitrate reduction
Nitrate reductase
Nitrate reductases are molybdoenzymes that reduce nitrate to nitrite .* Eukaryotic nitrate reductases are part of the sulfite oxidase family of molybdoenzymes....

 and nitrogen fixation
Nitrogen fixation
Nitrogen fixation is the natural process, either biological or abiotic, by which nitrogen in the atmosphere is converted into ammonia . This process is essential for life because fixed nitrogen is required to biosynthesize the basic building blocks of life, e.g., nucleotides for DNA and RNA and...

. A major source of iron to the oceans is dust from the Earth's desert
A desert is a landscape or region that receives an extremely low amount of precipitation, less than enough to support growth of most plants. Most deserts have an average annual precipitation of less than...

s, picked up and delivered by the wind as aeolian dust.

In regions of the ocean that are distant from deserts or that are not reached by dust-carrying winds (for example, the Southern
Southern Ocean
The Southern Ocean comprises the southernmost waters of the World Ocean, generally taken to be south of 60°S latitude and encircling Antarctica. It is usually regarded as the fourth-largest of the five principal oceanic divisions...

 and North Pacific oceans), the lack of iron can severely limit the amount of primary production that can occur. These areas are sometimes known as HNLC
HNLC stands for "high-nitrate, low-chlorophyll" - a term used in marine ecology to describe areas of the ocean where the number of phytoplankton are low and fairly constant in spite of high macro-nutrient concentrations...

 (High-Nutrient, Low-Chlorophyll) regions, because the scarcity of iron both limits phytoplankton growth and leaves a surplus of other nutrients. Some scientists have suggested introducing iron
Iron fertilization
Iron fertilization is the intentional introduction of iron to the upper ocean to stimulate a phytoplankton bloom. This is intended to enhance biological productivity, which can benefit the marine food chain and remove carbon dioxide from the atmosphere. Iron is a trace element necessary for...

 to these areas as a means of increasing primary productivity and sequestering carbon dioxide from the atmosphere.


The methods for measurement of primary production vary depending on whether gross vs net production is the desired measure, and whether terrestrial or aquatic systems are the focus. Gross production is almost always harder to measure than net, because of respiration, which is a continuous and ongoing process that consumes some of the products of primary production (i.e. sugars) before they can be accurately measured. Also, terrestrial ecosystems are generally more difficult because a substantial proportion of total productivity is shunted to below-ground organs and tissues, where it is logistically difficult to measure. Shallow water aquatic systems can also face this problem.

Scale also greatly affects measurement techniques. The rate of carbon assimilation in plant tissues, organs, whole plants, or plankton samples can be quantified by biochemically-based techniques, but these techniques are decidedly inappropriate for large scale terrestrial field situations. There, net primary production is almost always the desired variable, and estimation techniques involve various methods of estimating dry-weight biomass changes over time. Biomass estimates are often converted to an energy measure, such as kilocalories, by an empirical
The word empirical denotes information gained by means of observation or experimentation. Empirical data are data produced by an experiment or observation....

ly determined conversion factor.


In terrestrial ecosystems, researchers generally measure net primary production. Although its definition is straightforward, field measurements used to estimate productivity vary according to investigator and biome. Field estimates rarely account for below ground productivity, herbivory, decomposition
Decomposition is the process by which organic material is broken down into simpler forms of matter. The process is essential for recycling the finite matter that occupies physical space in the biome. Bodies of living organisms begin to decompose shortly after death...

, turnover, litterfall
Litterfall is the transport of leaves, bark, twigs and other forms of dead organic material and its constituent nutrients from the aerial parts of the biosphere to the top layer of soil, commonly known as the litter layer or O horizon.Litterfall has occupied the attention of ecologists at length...

, volatile organic compounds, root exudates, and allocation to symbiotic microorganisms. Biomass based NPP estimates result in underestimation of NPP due to incomplete accounting of these components. However, many field measurements correlate well to NPP. There are a number of comprehensive reviews of the field methods used to estimate NPP. Estimates of ecosystem respiration
Ecosystem respiration
Ecosystem respiration is the sum of all respiration occurring by the living organisms in a specific ecosystem.Ecosystem respiration is typically measured in the natural environment, such as a forest or grassland field, rather than in the laboratory...

, the total carbon dioxide produced by the ecosystem, can also be made with gas flux measurements.

The major unaccounted for pool is belowground productivity, especially production and turnover of roots. Belowground components of NPP are difficult to measure. BNPP is often estimated based on a ratio of ANPP:BNPP rather than direct measurements.


Most frequently, peak standing biomass is assumed to measure NPP. In systems with persistent standing litter, live biomass is commonly reported. Measures of peak biomass are more reliable in if the system is predominantly annuals. However, perennial measurements can be reliable if there was a synchronous phenology driven by a strong seasonal climate. These methods may underestimate ANPP in grasslands by as much as 2 (temperate
In geography, temperate or tepid latitudes of the globe lie between the tropics and the polar circles. The changes in these regions between summer and winter are generally relatively moderate, rather than extreme hot or cold...

) to 4 (tropical) fold. Repeated measures of standing live and dead biomass provide more accurate estimates of all grasslands, particularly those with large turnover, rapid decomposition, and interspecific variation in timing of peak biomass. Wetland
A wetland is an area of land whose soil is saturated with water either permanently or seasonally. Wetlands are categorised by their characteristic vegetation, which is adapted to these unique soil conditions....

 productivity (marshes and fens) is similarly measured. In Europe
Europe is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally 'divided' from Asia to its east by the watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...

, annual mowing makes the annual biomass increment of wetlands evident.


Methods used to measure forest productivity are more diverse than those of grasslands. Biomass increment based on stand specific allometry plus litterfall is considered a suitable although incomplete accounting of above-ground net primary production (ANPP). Field measurements used as a proxy for ANPP include annual litterfall, diameter or basal area increment (DBH
Diameter at breast height
Diameter at breast height, or DBH, is a standard method of expressing the diameter of the trunk or bole of a standing tree. DBH is one of the most common dendrometric measurements....

 or BAI), and volume increment.


In aquatic systems, primary production is typically measured using one of four main techniques :
  1. variations in oxygen concentration within a sealed bottle (developed by Gaarder and Gran in 1927)
  2. incorporation of inorganic carbon-14
    Carbon-14, 14C, or radiocarbon, is a radioactive isotope of carbon with a nucleus containing 6 protons and 8 neutrons. Its presence in organic materials is the basis of the radiocarbon dating method pioneered by Willard Libby and colleagues , to date archaeological, geological, and hydrogeological...

     (14C in the form of sodium bicarbonate
    Sodium bicarbonate
    Sodium bicarbonate or sodium hydrogen carbonate is the chemical compound with the formula Na HCO3. Sodium bicarbonate is a white solid that is crystalline but often appears as a fine powder. It has a slightly salty, alkaline taste resembling that of washing soda . The natural mineral form is...

    ) into organic matter
  3. Stable isotopes of Oxygen (16O, 18O and 17O)
  4. fluorescence kinetics (technique still a research topic)

The technique developed by Gaarder and Gran uses variations in the concentration of oxygen under different experimental conditions to infer gross primary production. Typically, three identical transparent vessels are filled with sample water and stoppered
Stopper (plug)
A bung is truncated cylindrical or conical closure to seal a container, such as a bottle, tube or barrel. Unlike a lid which encloses a container from the outside without displacing the inner volume, a bung is partially inserted inside the container to act as a seal...

. The first is analysed immediately and used to determine the initial oxygen concentration; usually this is done by performing a Winkler titration
Winkler test for dissolved oxygen
The Winkler test is used to determine the concentration of dissolved oxygen in water samples. Dissolved oxygen is widely used in water quality studies and routine operation of water reclamation facilities. An excess of manganese salt, iodide and hydroxide ions is added to a water sample causing...

. The other two vessels are incubated, one each in under light and darkened. After a fixed period of time, the experiment ends, and the oxygen concentration in both vessels is measured. As photosynthesis has not taken place in the dark vessel, it provides a measure of ecosystem respiration
Ecosystem respiration
Ecosystem respiration is the sum of all respiration occurring by the living organisms in a specific ecosystem.Ecosystem respiration is typically measured in the natural environment, such as a forest or grassland field, rather than in the laboratory...

. The light vessel permits both photosynthesis and respiration, so provides a measure of net photosynthesis (i.e. oxygen production via photosynthesis subtract oxygen consumption by respiration). Gross primary production is then obtained by adding oxygen consumption in the dark vessel to net oxygen production in the light vessel.

The technique of using 14C incorporation (added as labelled Na2CO3) to infer primary production is most commonly used today because it is sensitive, and can be used in all ocean environments. As 14C is radioactive
Radioactive decay
Radioactive decay is the process by which an atomic nucleus of an unstable atom loses energy by emitting ionizing particles . The emission is spontaneous, in that the atom decays without any physical interaction with another particle from outside the atom...

 (via beta decay
Beta decay
In nuclear physics, beta decay is a type of radioactive decay in which a beta particle is emitted from an atom. There are two types of beta decay: beta minus and beta plus. In the case of beta decay that produces an electron emission, it is referred to as beta minus , while in the case of a...

), it is relatively straightforward to measure its incorporation in organic material using devices such as scintillation counter
Scintillation counter
A scintillation counter measures ionizing radiation. The sensor, called a scintillator, consists of a transparent crystal, usually phosphor, plastic , or organic liquid that fluoresces when struck by ionizing radiation. A sensitive photomultiplier tube measures the light from the crystal...


Depending upon the incubation time chosen, net or gross primary production can be estimated. Gross primary production is best estimated using relatively short incubation times (1 hour or less), since the loss of incorporated 14C (by respiration and organic material excretion / exudation) will be more limited. Net primary production is the fraction of gross production remaining after these loss processes have consumed some of the fixed carbon.

Loss processes can range between 10-60% of incorporated 14C according to the incubation period, ambient environmental conditions (especially temperature) and the experimental species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

 used. Aside from those caused by the physiology of the experimental subject itself, potential losses due to the activity of consumers also need to be considered. This is particularly true in experiments making use of natural assemblages of microscopic autotrophs, where it is not possible to isolate them from their consumers.


As primary production in the biosphere
The biosphere is the global sum of all ecosystems. It can also be called the zone of life on Earth, a closed and self-regulating system...

 is an important part of the carbon cycle
Carbon cycle
The carbon cycle is the biogeochemical cycle by which carbon is exchanged among the biosphere, pedosphere, geosphere, hydrosphere, and atmosphere of the Earth...

, estimating it at the global scale is important in Earth system science
Earth science
Earth science is an all-embracing term for the sciences related to the planet Earth. It is arguably a special case in planetary science, the Earth being the only known life-bearing planet. There are both reductionist and holistic approaches to Earth sciences...

. However, quantifying primary production at this scale is difficult because of the range of habitat
Habitat (ecology)
A habitat is an ecological or environmental area that is inhabited by a particular species of animal, plant or other type of organism...

s on Earth, and because of the impact of weather
Weather is the state of the atmosphere, to the degree that it is hot or cold, wet or dry, calm or stormy, clear or cloudy. Most weather phenomena occur in the troposphere, just below the stratosphere. Weather refers, generally, to day-to-day temperature and precipitation activity, whereas climate...

 events (availability of sunlight, water) on its variability.

Using satellite
In the context of spaceflight, a satellite is an object which has been placed into orbit by human endeavour. Such objects are sometimes called artificial satellites to distinguish them from natural satellites such as the Moon....

-derived estimates of the Normalized Difference Vegetation Index
Normalized Difference Vegetation Index
The Normalized Difference Vegetation Index is a simple graphical indicator that can be used to analyze remote sensing measurements, typically but not necessarily from a space platform, and assess whether the target being observed contains live green vegetation or not.-Brief history:The exploration...

 (NDVI) for terrestrial habitats and sea-surface chlorophyll
Chlorophyll is a green pigment found in almost all plants, algae, and cyanobacteria. Its name is derived from the Greek words χλωρος, chloros and φύλλον, phyllon . Chlorophyll is an extremely important biomolecule, critical in photosynthesis, which allows plants to obtain energy from light...

 for the oceans, it is estimated that the total (photoautotrophic) primary production for the Earth was 104.9 Gt
The tonne, known as the metric ton in the US , often put pleonastically as "metric tonne" to avoid confusion with ton, is a metric system unit of mass equal to 1000 kilograms. The tonne is not an International System of Units unit, but is accepted for use with the SI...

 C yr−1. Of this, 56.4 Gt C yr−1 (53.8%), was the product of terrestrial organisms, while the remaining 48.5 Gt C yr−1, was accounted for by oceanic production.

In area
Area is a quantity that expresses the extent of a two-dimensional surface or shape in the plane. Area can be understood as the amount of material with a given thickness that would be necessary to fashion a model of the shape, or the amount of paint necessary to cover the surface with a single coat...

l terms, it was estimated that land production was approximately 426 g C m−2 yr−1 (excluding areas with permanent ice cover), while that for the oceans was 140 g C m−2 yr−1. Another significant difference between the land and the oceans lies in their standing stocks - while accounting for almost half of total production, oceanic autotrophs only account for about 0.2% of the total biomass.

Human impact and appropriation

Extensive human land use
Land use
Land use is the human use of land. Land use involves the management and modification of natural environment or wilderness into built environment such as fields, pastures, and settlements. It has also been defined as "the arrangements, activities and inputs people undertake in a certain land cover...

 results in various levels of impact on actual NPP (NPPact). In some regions, such as the Nile
The Nile is a major north-flowing river in North Africa, generally regarded as the longest river in the world. It is long. It runs through the ten countries of Sudan, South Sudan, Burundi, Rwanda, Democratic Republic of the Congo, Tanzania, Kenya, Ethiopia, Uganda and Egypt.The Nile has two major...

 valley, irrigation
Irrigation may be defined as the science of artificial application of water to the land or soil. It is used to assist in the growing of agricultural crops, maintenance of landscapes, and revegetation of disturbed soils in dry areas and during periods of inadequate rainfall...

 has resulted in a considerable increase in primary production. However, these regions are exceptions to the rule, and in general there is a NPP reduction due to land changes (ΔNPPLC) of 9.6% across global land-mass. In addition to this, end consumption by people raises the total human appropriation of net primary production (HANPP) to 23.8% of potential vegetation (NPP0). It is estimated that, in 2000, 34% of the Earth's ice-free land area (12% cropland
Arable land
In geography and agriculture, arable land is land that can be used for growing crops. It includes all land under temporary crops , temporary meadows for mowing or pasture, land under market and kitchen gardens and land temporarily fallow...

; 22% pasture
Pasture is land used for grazing. Pasture lands in the narrow sense are enclosed tracts of farmland, grazed by domesticated livestock, such as horses, cattle, sheep or swine. The vegetation of tended pasture, forage, consists mainly of grasses, with an interspersion of legumes and other forbs...

) was devoted to human agriculture. This disproportionate amount reduces the energy available to other species, having a marked impact on biodiversity
Biodiversity is the degree of variation of life forms within a given ecosystem, biome, or an entire planet. Biodiversity is a measure of the health of ecosystems. Biodiversity is in part a function of climate. In terrestrial habitats, tropical regions are typically rich whereas polar regions...

, flows of carbon, water and energy, and ecosystem services
Ecosystem services
Humankind benefits from a multitude of resources and processes that are supplied by natural ecosystems. Collectively, these benefits are known as ecosystem services and include products like clean drinking water and processes such as the decomposition of wastes...

, and scientists have questioned how large this fraction can be before these services begin to break down.
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