Primary production

804:, 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. 1753:/Ar can even be measured continuously at sea using equilibrator inlet mass spectrometry (EIMS) or a membrane inlet mass spectrometry (MIMS). However, if results relevant to the carbon cycle are desired, it is probably better to rely on methods based on carbon (and not oxygen) isotopes. It is important to notice that the method based on carbon stable isotopes is not simply an adaptation of the classic C method, but an entirely different approach that does not suffer from the problem of lack of account of carbon recycling during photosynthesis. 501: 1036: 754: 27: 694:(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 599: 492:

abundant sunlight, warmth, and rainfall. However, even in the tropics, there are variations in productivity over the course of the year. For example, the Amazon basin exhibits especially high productivity from roughly August through October - the period of the area's dry season. Because the trees have access to a plentiful supply of ground water that builds up in the rainy season, they grow better when the rainy skies clear and allow more sunlight to reach the forest.

1613: 2016:"Gross primary production" (GPP) refers to the total rate of organic carbon production by autotrophs, while "respiration" refers to the energy-yielding oxidation of organic carbon back to carbon dioxide. "Net primary production"(NPP) is GPP minus the autotrophs' own rate of respiration; it is thus the rate at which the full metabolism of phytoplankton produces biomass. "Secondary production" (SP) typically refers to the growth rate of heterotrophic biomass. 645: 159: 871:

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.

991: 891: 944: 878:, 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 1866:

introduced it at the inaugural conference of the European Society for Ecological Economics. HANPP has since been further developed and widely applied in research on ecological economics and in policy analysis for sustainability. HANPP represents a proxy of the human impact on nature and can be applied to different geographical and global scales.

430:(the latter covaries to an extent with light, specifically photosynthetically active radiation (PAR), 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 PAR) are limiting, such as 379: 1703:. 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. 1921:,. Scientists have questioned how large this fraction can be before these services break down. Reductions in NPP are also expected in the ocean as a result of ongoing climate change, potentially impacting marine ecosystems (~10% of global biodiversity) and goods and services (1-5% of global total) that the oceans provide. 1823:

terms, it was estimated that land production was approximately 426 g C m yr (excluding areas with permanent ice cover), while that for the oceans was 140 g C m yr. Another significant difference between the land and the oceans lies in their standing stocks - while accounting for almost half of total

1737:

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

1865:

Human societies are part of the Earth's NPP cycle but disproportionately influence it. In 1996, Josep Garí designed a new indicator of sustainable development based precisely on the estimation of the human appropriation of NPP: he coined it "HANPP" (Human Appropriation of Net Primary Production) and

1729:

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 C (by respiration and organic material excretion / exudation) will be more

663:(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 387:

This animation shows Earth's monthly terrestrial net primary productivity from 2000 to 2013. Values range from near 0 grams of carbon per square meter per day (tan) to 6.5 grams per square meter per day (dark green). A negative value means decomposition or respiration overpowered carbon absorption;

927:

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

388:

more carbon was released to the atmosphere than the plants took in. In mid-latitudes, productivity obviously interacts with seasonal change, with productivity peaking in each hemisphere's summer. The data come from the Moderate Resolution Imaging Spectroradiometer (MODIS) on NASA's Terra satellite.

636:, 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. 965:

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 if the system is predominantly annuals. However, perennial measurements could be reliable if there were a

902:

In terrestrial ecosystems, researchers generally measure net primary production (NPP). 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,

870:

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

491:

As shown in the animation, the boreal forests of Canada and Russia experience high productivity in June and July and then a slow decline through fall and winter. Year-round, tropical forests in South America, Africa, Southeast Asia, and Indonesia have high productivity, not surprising with the

369:

Both gross and net primary production are typically expressed in units of mass per unit area per unit time interval. In terrestrial ecosystems, mass of carbon per unit area per year (g C m yr) is most often used as the unit of measurement. Note that a distinction is sometimes drawn between

915:

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

3077:

Crockford, Peter W.; Kunzmann, Marcus; Bekker, Andrey; Hayles, Justin; Bao, Huiming; Halverson, Galen P.; Peng, Yongbo; Bui, Thi H.; Cox, Grant M.; Gibson, Timothy M.; Wörndle, Sarah; Rainbird, Robert; Lepland, Aivo; Swanson-Hysell, Nicholas L.; Master, Sharad (2019-05-20).

1699:. 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 1832:

Present day primary productivity can be estimated through a variety of methodologies including ship-board measurements, satellites and terrestrial observatories. Historical estimates have relied on biogeochemical models and geochemical proxies. One example is using

468:. These structures 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 383: 1690:

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

380: 2423:

Reichstein, Markus; Falge, Eva; Baldocchi, Dennis; Papale, Dario; Aubinet, Marc; Berbigier, Paul; et al. (2005). "On the separation of net ecosystem exchange into assimilation and ecosystem respiration: review and improved algorithm".

974:) 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. 1006:

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

382: 370:"production" and "productivity", with the former the quantity of material produced (g C m), the latter the rate at which it is produced (g C m yr), but these terms are more typically used interchangeably. 2474:

Marra, J. (2002), pp. 78-108. In: Williams, P. J. leB., Thomas, D. N., Reynolds, C. S. (Eds.), Phytoplankton Productivity:Carbon Assimilation in Marine and Freshwater Ecosystems. Blackwell, Oxford, UK

366:

produce net useful chemical energy. Net primary production is available to be directed toward growth and reproduction of primary producers. As such it is available for consumption by herbivores.

2566: 1745:/Ar ratios have the advantage of providing estimates of respiration rates in the light without the need of incubations in the dark. Among them, the method of the triple oxygen isotopes and O 3038:

Crockford, Peter W.; Hayles, Justin A.; Bao, Huiming; Planavsky, Noah J.; Bekker, Andrey; Fralick, Philip W.; Halverson, Galen P.; Bui, Thi Hao; Peng, Yongbo; Wing, Boswell A. (July 2018).

1808:

measurements of net ecosystem exchange (see above) to regional and global values using spatial details of different predictor variables, such as climate variables and remotely sensed

701:

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

2940:

Prakash Babua, C.; Brumsack, H.-J.; Schnetger, B.; Böttcher, M.E. (2002). "Barium as a productivity proxy in continental margin sediments: a study from the eastern Arabian Sea".

1733:

Loss processes can range between 10 and 60% of incorporated C according to the incubation period, ambient environmental conditions (especially temperature) and the experimental

858:(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 1637: 2769:

Kaiser, J.; M. K. Reuer; B. Barnett; M.L. Bender (2005). "Marine productivity estimates from continuous O-2/Ar ratio measurements by membrane inlet mass spectrometry".

381: 2295:

Clark, D.A.; Brown, S.; Kicklighter, D.W.; Chambers, J.Q.; Thomlinson, J.R.; Ni, J. (2001). "Measuring net primary production in forests: Concepts and field methods".

874:

Scale also greatly affects measurement techniques. The rate of carbon assimilation in plant tissues, organs, whole plants, or plankton samples can be quantified by

606:

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

935:. During night, this technique measures all components of ecosystem respiration. This respiration is scaled to day-time values and further subtracted from NEE. 484:

adaptations to increase water-use efficiency and allow increased primary production to take place under conditions that would normally limit carbon fixation by

1801:

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

875: 464:

to growth regions, and also cools the plant. Diffusion of water vapour out of a leaf, the force that drives transpiration, is regulated by structures known as

1684: 999: 3163: 2130:

The deep chlorophyll maximum (DCM) occurs at the contact where there is adequate light for photosynthesis and yet significant nutrient supply from below.

725:

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

698:. As long as there are adequate nutrients available, net primary production occurs whenever the mixed layer is shallower than the critical depth. 1730:

limited. Net primary production is the fraction of gross production remaining after these loss processes have consumed some of the fixed carbon.

3354: 3245: 800:

of deeper water. This is exacerbated where summertime solar heating and reduced winds increases vertical stratification and leads to a strong

1913:) was devoted to human agriculture. This disproportionate amount reduces the energy available to other species, having a marked impact on 522:

In a reversal of the pattern on land, in the oceans, almost all photosynthesis is performed by algae, with a small fraction contributed by

2107: 1849:. Together these records suggest large shifts in primary production throughout Earth's past with notable rises associated with Earth's 1714:) to infer primary production is most commonly used today because it is sensitive, and can be used in all ocean environments. As C is 1123: 1408: 1081: 2601:

Luz and Barkan, B; Barkan, E (2000). "Assessment of oceanic productivity with the triple-isotope composition of dissolved oxygen".

1695:. The first is analysed immediately and used to determine the initial oxygen concentration; usually this is done by performing a 854:

oceans), the lack of iron can severely limit the amount of primary production that can occur. These areas are sometimes known as

3239:

Haberl, H.; Erb, K.H.; Krausmann, F.; Gaube, V.; Bondeau, A.; Plutzar, C.; Gingrich, S.; Lucht, W.; Fischer-Kowalski, M. (2007).

1581: 1108: 1786: 1644: 1462: 966:

synchronous phenology driven by a strong seasonal climate. These methods may underestimate ANPP in grasslands by as much as 2 (

334:, that primary producers create in a given length of time. Some fraction of this fixed energy is used by primary producers for 2407: 1586: 1841:

concentrations in marine sediments rise in line with carbon export production at the surface. Another example is using the

86:

on Earth relies directly or indirectly on primary production. The organisms responsible for primary production are known as

2196:

Martin, J. H.; Fitzwater, S. E. (1988). "Iron-deficiency limits phytoplankton growth in the Northeast Pacific Subarctic".

931:

Gross primary production can be estimated from measurements of net ecosystem exchange (NEE) of carbon dioxide made by the

38:

biomass, it is only a rough indicator of primary-production potential, and not an actual estimate of it. Provided by the

2730:

Cassar, N.; B.A. Barnett; M.L. Bender; J. Kaiser; R.C. Hamme; B. Tilbrooke (2009). "Continuous high-frequency dissolved O

1051: 393: 772:

Mixing also plays an important role in the limitation of primary production by nutrients. Inorganic nutrients, such as

1897:) of 9.6% across global land-mass. In addition to this, end consumption by people raises the total HANPP to 23.8% of 846:

In regions of the ocean that are distant from deserts or that are not reached by dust-carrying winds (for example, the

558:, and a diverse group of unicellular groups. Vascular plants are also represented in the ocean by groups such as the 1696: 796:, dead or fecal material), nutrients are constantly lost from the photic zone, and are only replenished by mixing or 632:

it at lower temperatures. However, the availability of light, the source of energy for photosynthesis, and mineral

1960: 2564:

Bender, Michael; et al. (1987). "A Comparison of 4 Methods for Determining Planktonic Community Production".

1889:

has resulted in a considerable increase in primary production, in most of the Planet, there is a notable trend of

1816:

led to a terrestrial gross primary production of 123±8 Gt carbon (NOT carbon dioxide) per year during 1998-2005

862:

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

279:. These relatively simple molecules may be then used to further synthesise more complicated molecules, including 3194: 2308: 526:

and other groups. Algae encompass a diverse range of organisms, ranging from single floating cells to attached

418:

of vascular plants, non-vascular plants likely played a more significant role. Primary production on land is a

2537:

Steeman-Nielsen, E. (1952). "The use of radioactive carbon (C14) for measuring organic production in the sea".

706: 1978:

Amthor, J.S. and Baldocchi, D.D. (2001). Terrestrial Higher Plant Respiration and Net Primary Production. In

812:

Another factor relatively recently discovered to play a significant role in oceanic primary production is the

737:, light may only vary slightly across the year, and mixing may only occur episodically, such as during large 3241:"Quantifying and mapping the human appropriation of net primary production in Earth's terrestrial ecosystems" 1793:

for the oceans, it is estimated that the total (photoautotrophic) primary production for the Earth was 104.9

615: 82:, which uses the oxidation or reduction of inorganic chemical compounds as its source of energy. Almost all 20: 1722:), it is relatively straightforward to measure its incorporation in organic material using devices such as 1516: 1511: 1501: 1181: 565:

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

469: 1993: 190:

O). The following two equations are simplified representations of photosynthesis (top) and (one form of)

2679:

Craig and Hayward (1987). "Oxygen supersaturations in the ocean: biological vs. physical contributions".

47: 2484:

Steeman-Nielsen, E. (1951). "Measurement of production of organic matter in sea by means of carbon-14".

2031: 678:

Net photosynthesis in the water column is determined by the interaction between the photic zone and the

1571: 1469: 1175: 1169: 908: 3079: 1576: 1196: 1008: 517: 2791: 2147:"Biotic and Human Vulnerability to Projected Changes in Ocean Biogeochemistry over the 21st Century" 2989: 1551: 1479: 1474: 1457: 1302: 1163: 1061: 675:

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

2874:

Beer, C.; Reichstein, M.; Tomelleri, E.; Ciais, P.; Jung, M.; Carvalhais, N.; et al. (2010).

3211:

Garí, J.A. (1996). "HANPP calculated from land cover as indicator of ecological sustainability".

2875: 1661:

variations in oxygen concentration within a sealed bottle (developed by Gaarder and Gran in 1927)

1561: 1076: 820: 339: 3120:

Crockford, Peter W.; Bar On, Yinon M.; Ward, Luce M.; Milo, Ron; Halevy, Itay (November 2023).

2786: 1945: 1940: 1657:

In aquatic systems, primary production is typically measured using one of six main techniques:

1381: 1157: 1118: 1086: 419: 3301:"Farming the planet: 1. Geographic distribution of global agricultural lands in the year 2000" 1773:. However, quantifying primary production at this scale is difficult because of the range of 595:, but the vast majority of free-floating production takes place within microscopic organisms. 1850: 1723: 1700: 1630: 1617: 1566: 1297: 917: 300: 284: 101: 3423: 3362: 3312: 3253: 3004: 2949: 2890: 2876:"Terrestrial Gross Carbon Dioxide Uptake: Global Distribution and Covariation with Climate" 2840: 2778: 2688: 2610: 2575: 2495: 2433: 2356: 2256: 2205: 1493: 1395: 1151: 1056: 690:

energy at the ocean's surface homogenises the water column vertically until the turbulence

534:

are important photosynthetizers in both oceanic and terrestrial ecosystems, and while some

335: 296: 124: 602:

Differences in relative photosynthesis between plankton species under different irradiance

8: 2239:

Cooper, D.J.; Watson, A.J.; Nightingale, P.D. (1996). "Large decrease in ocean—surface CO

1442: 1258: 1113: 668: 445:. The latter process (which is responsible for about 90% of water use) is driven by the 3366: 3316: 3257: 3008: 2953: 2894: 2844: 2782: 2692: 2614: 2579: 2499: 2437: 2360: 2260: 2209: 3385: 3349: 3330: 3276: 3240: 3186: 3020: 2922: 2804: 2712: 2661: 2519: 2457: 2392: 2372: 2320: 2272: 2221: 2173: 2146: 1918: 1842: 1749:/Ar have the additional advantage of not needing incubations in closed containers and O 1669: 1521: 1289: 1273: 1268: 859: 629: 582: 485: 473: 403: 2961: 2445: 2083: 2058: 3390: 3334: 3281: 3141: 3099: 3059: 3024: 2926: 2914: 2906: 2856: 2831: 2827:"Primary production of the Biosphere: Integrating Terrestrial and Oceanic Components" 2751: 2704: 2626: 2511: 2449: 2403: 2368: 2312: 2178: 2088: 1935: 1774: 1715: 1452: 1231: 1066: 1002:

are more diverse than those of grasslands. Biomass increment based on stand specific

951: 840: 832: 828: 785: 762: 331: 109: 88: 31: 3039: 2808: 2716: 2665: 2644:

Carvalho and Eyre (2012). "Measurement of planktonic CO2 respiration in the light".

2461: 2376: 2324: 311:, then transfers these organic molecules (and the energy stored within them) up the 3418: 3380: 3370: 3320: 3271: 3261: 3216: 3178: 3133: 3091: 3051: 3012: 2957: 2898: 2848: 2796: 2743: 2696: 2653: 2618: 2583: 2546: 2523: 2503: 2486: 2441: 2364: 2304: 2276: 2264: 2225: 2213: 2168: 2158: 2078: 2074: 2070: 1950: 1418: 1312: 1307: 765: 171: 67: 3095: 2622: 2345:"Estimating net primary productivity from grassland biomass dynamics measurements" 714: 3413: 3040:"Triple oxygen isotope evidence for limited mid-Proterozoic primary productivity" 2852: 2824: 2700: 2399: 2163: 1930: 1824:

production, oceanic autotrophs only account for about 0.2% of the total biomass.

1813: 1805: 1546: 1413: 1346: 1334: 1263: 1137: 932: 921: 523: 500: 441:

Water is "consumed" in plants by the processes of photosynthesis (see above) and

411: 136: 2826: 2550: 1854: 1692: 1556: 1526: 1506: 1447: 1361: 1244: 1239: 1100: 847: 695: 399: 342:"). The remaining fixed energy (i.e., mass of photosynthate) is referred to as 191: 175: 79: 75: 71: 3161: 3137: 3055: 2587: 1905:). It is estimated that, in 2000, 34% of the Earth's ice-free land area (12% 115:

predominate in this role. Ecologists distinguish primary production as either

3407: 3145: 3103: 3063: 2910: 2453: 2316: 2294: 1798: 1770: 948: 813: 734: 622: 570: 566: 442: 292: 3375: 3266: 3121: 2902: 2657: 1035: 920:, the total carbon dioxide produced by the ecosystem, can also be made with 3394: 3298: 3285: 2918: 2755: 2708: 2630: 2515: 2182: 2092: 1914: 1766: 1339: 1253: 1207: 1145: 1027: 781: 753: 546:

are significant contributors to primary production in the ocean, including

435: 260: 163: 26: 16:

Synthesis of organic compounds from carbon dioxide by biological organisms

3080:"Claypool continued: Extending the isotopic record of sedimentary sulfate" 2860: 598: 3325: 3300: 2800: 1906: 1790: 1403: 1376: 1371: 1366: 1356: 1326: 1212: 1071: 955: 801: 758: 691: 679: 664: 660: 649: 586: 542:, none are known to utilise oxygen-evolving photosynthesis. A number of 446: 427: 304: 3220: 2990:"Barium in Deep-Sea Sediment: A Geochemical Proxy for Paleoproductivity" 1869:

The extensive degree of human use of the Planet's resources, mostly via

982:, annual mowing makes the annual biomass increment of wetlands evident. 3347: 3190: 1886: 1719: 1351: 904: 710: 683: 672: 653: 644: 611: 551: 547: 539: 477: 148: 97: 78:, which uses light as its source of energy, but it also occurs through 3213:

Proceedings of the International Conference: Ecology, Society, Economy

3016: 2939: 2825:

Field, C.B.; Behrenfeld, M.J.; Randerson, J.T.; Falkowski, P. (1998).

2768: 2747: 2729: 330:(GPP) is the amount of chemical energy, typically expressed as carbon 322: 3238: 2507: 2268: 2217: 1782: 1762: 1665: 1003: 967: 912: 879: 797: 777: 742: 718: 633: 591: 543: 423: 415: 363: 362:

Net primary production is the rate at which all the autotrophs in an

338:

and maintenance of existing tissues (i.e., "growth respiration" and "

152: 93: 35: 3182: 2344: 1781:

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

170:

Regardless of its source, this energy is used to synthesize complex

3162:

Vitousek, P.M.; Ehrlich, P.R.; Ehrlich, A.H.; Matson, P.A. (1986).

2238: 1870: 971: 793: 722: 607: 578: 559: 555: 531: 527: 509: 457: 312: 144: 140: 34:

abundance, from September 1997 to August 2000. As an estimate of

1910: 1846: 1778: 1734: 1434: 1217: 975: 851: 835:. A major source of iron to the oceans is dust from the Earth's 789: 773: 726: 626: 574: 535: 481: 454: 280: 272: 253: 59: 39: 158: 2974:

E.D. Goldberg, G.O.S. Arrhenius Chemistry of pelagic sediments

1838: 1834: 1596: 979: 959: 836: 824: 702: 505: 453:

of plants. Transpiration allows plants to transport water and

431: 398:

On the land, almost all primary production is now performed by

308: 288: 51: 3299:

Ramankutty, N.; Evan, A.T.; Monfreda, C.; Foley, J.A. (2008).

2422: 2342: 625:

of seawater buffers temperature changes, and the formation of

3076: 1809: 1794: 1591: 738: 465: 316: 276: 257: 183: 112: 105: 3348:

Foley, J.A.; Monfreda, C.; Ramankutty, N.; Zaks, D. (2007).

2873: 1706:

The technique of using C incorporation (added as labelled Na

990: 890: 621:), ranges less widely in the ocean than on land because the 573:. Larger autotrophs, such as the seagrasses and macroalgae ( 3037: 2734:/Ar measurements by equilibrator inlet mass spectrometry". 2309:

10.1890/1051-0761(2001)011[0356:MNPPIF]2.0.CO;2

1882: 1820: 855: 816: 730: 687: 530:. They include photoautotrophs from a variety of groups. 461: 450: 407: 83: 43: 978:

productivity (marshes and fens) is similarly measured. In

3115: 3113: 943: 895: 147:

but a minute fraction of primary production is driven by

2563: 1678:

fluorescence kinetics (technique still a research topic)

123:, the former accounting for losses to processes such as 3164:"Human appropriation of the products of photosynthesis" 3119: 3110: 2056: 2057:

Schäfer G, Engelhard M, Müller V (1 September 1999).

1853:(approximately 2.4 to 2.0 billion years ago) and the 713:), although wind magnitudes additionally have strong 2600: 2389: 1769:, estimating it at the global scale is important in 585:

to the underlying substrate but still be within the

2416: 2343:Scurlock, J.M.O.; Johnson, K.; Olson, R.J. (2002). 2099: 1797:

of carbon per year (Pg C yr; equivalent to the non-

323:

Gross primary production and net primary production

2867: 2391: 1985: 610:can be). Similarly, temperature, while affecting 2678: 2643: 1860: 1804:Scaling ecosystem-level GPP estimations based on 581:zone and adjacent shallow waters, where they can 3405: 2988:Dymond, Jack; Suess, Erwin; Lyle, Mitch (1992). 1789:(NDVI) for terrestrial habitats and sea-surface 788:their cells and cellular machinery. Because of 74:. It principally occurs through the process of 2987: 2820: 2818: 2536: 2483: 2195: 1857:(approximately 1.0 to 0.54 billion years ago). 3122:"The geologic history of primary productivity" 402:, with a small fraction coming from algae and 3157: 3155: 2032:"Net Primary Productivity : Global Maps" 1827: 1638: 898:tree; a typical modern, terrestrial autotroph 2815: 3234: 3232: 3230: 659:The sunlit zone of the ocean is called the 3152: 2108:"The Biological Productivity of the Ocean" 2105: 2026: 2024: 1994:"The Biological Productivity of the Ocean" 1991: 1917:, flows of carbon, water, and energy, and 1741:The methods based on stable isotopes and O 1645: 1631: 3384: 3374: 3324: 3275: 3265: 2790: 2172: 2162: 2082: 1881:). Although in some regions, such as the 1873:, results in various levels of impact on 839:, picked up and delivered by the wind as 792:sinking of particulate material (such as 373: 174:from simpler inorganic compounds such as 3227: 2338: 2336: 2334: 989: 942: 889: 757:Annual mean sea surface nitrate for the 752: 643: 597: 499: 488:plants (the majority of plant species). 377: 157: 135:Primary production is the production of 25: 2290: 2288: 2286: 2232: 2021: 1777:on Earth, and because of the impact of 1582:Territorialisation of carbon governance 717:. Consequently, primary production in 422:of many factors, but principally local 151:organisms using the chemical energy of 3406: 2140: 2138: 2050: 1787:Normalized Difference Vegetation Index 1675:Stable isotopes of Oxygen (O, O and O) 303:. Consumption of primary producers by 2394:Primary Productivity of the Biosphere 2331: 1587:Total Carbon Column Observing Network 495: 143:. The main source of this energy is 3210: 2283: 2144: 2646:Limnology and Oceanography: Methods 2390:Leith, H.; Whittaker, R.H. (1975). 2135: 1681:Stable isotopes of Carbon (C and C) 911:, root exudates, and allocation to 784:are necessary for phytoplankton to 394:Terrestrial biological carbon cycle 13: 252:In both cases, the end point is a 14: 3435: 2446:10.1111/j.1365-2486.2005.001002.x 2106:Sigman, D.M.; Hain, M.P. (2012). 1992:Sigman, D.M.; Hain, M.P. (2012). 1891:NPP reduction due to land changes 589:. There are exceptions, such as 3350:"Our share of the planetary pie" 2369:10.1046/j.1365-2486.2002.00512.x 1961:Lists of organisms by population 1612: 1611: 1034: 577:) are generally confined to the 3341: 3292: 3204: 3070: 3031: 2981: 2968: 2933: 2762: 2723: 2672: 2637: 2594: 2557: 2530: 2477: 2468: 2383: 1980:Terrestrial Global Productivity 1011:or BAI), and volume increment. 139:in organic compounds by living 30:Global oceanic and terrestrial 2189: 2145:Mora, C.; et al. (2013). 2075:10.1128/MMBR.63.3.570-620.1999 2059:"Bioenergetics of the Archaea" 1972: 1861:Human impact and appropriation 1547:Climate reconstruction proxies 885: 882:determined conversion factor. 876:biochemically based techniques 865: 827:involved in processes such as 271:, typically molecules such as 21:Primary production (economics) 1: 3355:Proc. Natl. Acad. Sci. U.S.A. 3246:Proc. Natl. Acad. Sci. U.S.A. 3096:10.1016/j.chemgeo.2019.02.030 2962:10.1016/S0025-3227(01)00286-9 2623:10.1126/science.288.5473.2028 1966: 1761:As primary production in the 938: 667:down the water column by its 3305:Global Biogeochemical Cycles 2853:10.1126/science.281.5374.237 2701:10.1126/science.235.4785.199 2164:10.1371/journal.pbio.1001682 1765:is an important part of the 1517:Carbonate compensation depth 1182:Particulate inorganic carbon 748: 470:Crassulacean acid metabolism 70:from atmospheric or aqueous 7: 1924: 1738:them from their consumers. 1664:incorporation of inorganic 733:in the middle of the major 130: 96:, and form the base of the 48:Goddard Space Flight Center 10: 3440: 2567:Limnology and Oceanography 2115:Nature Education Knowledge 2001:Nature Education Knowledge 1828:Present and Past Estimates 1785:-derived estimates of the 1572:Carbon capture and storage 1176:Particulate organic carbon 1170:Dissolved inorganic carbon 1014: 985: 909:volatile organic compounds 515: 391: 18: 3138:10.1016/j.cub.2023.09.040 3056:10.1038/s41586-018-0349-y 2588:10.4319/lo.1987.32.5.1085 2539:J. Cons. Int. Explor. Mer 2063:Microbiol. Mol. Biol. Rev 2036:earthobservatory.nasa.gov 1756: 1577:Carbon cycle re-balancing 933:eddy covariance technique 652:; an example of attached 518:Marine primary production 2978:, 13 (1958), pp. 153-212 2976:Geochim. Cosmochim. Acta 2551:10.1093/icesjms/18.2.117 2243:fugacity in response to 1552:Carbon-to-nitrogen ratio 1512:Carbonate–silicate cycle 1480:Carbon dioxide clathrate 1475:Clathrate gun hypothesis 1303:Net ecosystem production 1164:Dissolved organic carbon 998:Methods used to measure 639: 354:NPP = GPP - respiration 328:Gross primary production 3376:10.1073/pnas.0705190104 3267:10.1073/pnas.0704243104 2903:10.1126/science.1184984 2658:10.4319/lom.2012.10.167 2297:Ecological Applications 1982:, Academic Press, 33-59 1562:Deep Carbon Observatory 1022:Part of a series on the 807: 340:maintenance respiration 1843:triple oxygen isotopes 1724:scintillation counters 1382:Continental shelf pump 1158:Total inorganic carbon 1124:Satellite measurements 995: 962: 899: 819:. This is used as a 769: 656: 603: 513: 389: 374:Terrestrial production 344:net primary production 167: 102:terrestrial ecoregions 55: 2426:Global Change Biology 2349:Global Change Biology 2247:iron fertilization". 1851:Great Oxidation Event 1701:ecosystem respiration 1672:) into organic matter 1567:Global Carbon Project 1298:Ecosystem respiration 993: 946: 922:gas flux measurements 918:ecosystem respiration 893: 756: 729:regions, such as the 647: 601: 567:microscopic organisms 503: 386: 315:, fueling all of the 285:complex carbohydrates 161: 29: 3326:10.1029/2007GB002952 3132:(21): 4741–4750.e5. 2801:10.1029/2005GL023459 1899:potential vegetation 1771:Earth system science 1396:Carbon sequestration 1152:Total organic carbon 721:regions such as the 336:cellular respiration 125:cellular respiration 66:is the synthesis of 19:For other uses, see 3367:2007PNAS..10412585F 3361:(31): 12585–12586. 3317:2008GBioC..22.1003R 3258:2007PNAS..10412942H 3252:(31): 12942–12947. 3009:1992PalOc...7..163D 2954:2002MGeol.184..189P 2895:2010Sci...329..834B 2845:1998Sci...281..237F 2783:2005GeoRL..3219605K 2693:1987Sci...235..199C 2615:2000Sci...288.2028L 2609:(5473): 2028–2031. 2580:1987LimOc..32.1085B 2500:1951Natur.167..684N 2438:2005GCBio..11.1424R 2361:2002GCBio...8..736S 2261:1996Natur.383..511C 2210:1988Natur.331..341M 1685:Oxygen/Argon Ratios 1443:Atmospheric methane 1409:Soil carbon storage 1259:Reverse Krebs cycle 1114:Ocean acidification 1000:forest productivity 404:non-vascular plants 319:'s living systems. 307:organisms, such as 104:, these are mainly 2771:Geophys. Res. Lett 1919:ecosystem services 1670:sodium bicarbonate 1668:(C in the form of 1522:Great Calcite Belt 1470:Aerobic production 1290:Carbon respiration 1232:Metabolic pathways 1192:Primary production 996: 963: 900: 770: 715:spatial components 657: 604: 514: 496:Oceanic production 449:of water from the 390: 168: 127:, the latter not. 110:aquatic ecoregions 64:primary production 56: 3050:(7715): 613–616. 3017:10.1029/92PA00181 2889:(5993): 834–838. 2839:(5374): 237–240. 2748:10.1021/ac802300u 2687:(4785): 199–202. 2494:(4252): 684–685. 2409:978-0-387-07083-4 2255:(6600): 511–513. 2204:(6154): 341–343. 1936:Biomass (ecology) 1697:Winkler titration 1655: 1654: 1453:Methane emissions 1109:In the atmosphere 952:tallgrass prairie 833:nitrogen fixation 829:nitrate reduction 763:World Ocean Atlas 761:. Data from the 384: 172:organic molecules 166:of photosynthesis 89:primary producers 68:organic compounds 3431: 3399: 3398: 3388: 3378: 3345: 3339: 3338: 3328: 3296: 3290: 3289: 3279: 3269: 3236: 3225: 3224: 3208: 3202: 3201: 3199: 3193:. Archived from 3168: 3159: 3150: 3149: 3117: 3108: 3107: 3084:Chemical Geology 3074: 3068: 3067: 3035: 3029: 3028: 2997:Paleoceanography 2994: 2985: 2979: 2972: 2966: 2965: 2948:(3–4): 189–206. 2937: 2931: 2930: 2880: 2871: 2865: 2864: 2822: 2813: 2812: 2794: 2766: 2760: 2759: 2742:(5): 1855–1864. 2727: 2721: 2720: 2676: 2670: 2669: 2641: 2635: 2634: 2598: 2592: 2591: 2574:(5): 1085–1098. 2561: 2555: 2554: 2534: 2528: 2527: 2508:10.1038/167684b0 2481: 2475: 2472: 2466: 2465: 2432:(9): 1424–1439. 2420: 2414: 2413: 2397: 2387: 2381: 2380: 2340: 2329: 2328: 2292: 2281: 2280: 2269:10.1038/383511a0 2236: 2230: 2229: 2218:10.1038/331341a0 2193: 2187: 2186: 2176: 2166: 2157:(10): e1001682. 2142: 2133: 2132: 2127: 2126: 2112: 2103: 2097: 2096: 2086: 2054: 2048: 2047: 2045: 2043: 2028: 2019: 2018: 2013: 2012: 1998: 1989: 1983: 1976: 1951:Hydrogen sulfide 1647: 1640: 1633: 1620: 1615: 1614: 1419:pelagic sediment 1313:Soil respiration 1308:Photorespiration 1038: 1019: 1018: 958:of northeastern 860:introducing iron 684:Turbulent mixing 508:; an example of 385: 3439: 3438: 3434: 3433: 3432: 3430: 3429: 3428: 3404: 3403: 3402: 3346: 3342: 3297: 3293: 3237: 3228: 3209: 3205: 3197: 3183:10.2307/1310258 3166: 3160: 3153: 3126:Current Biology 3118: 3111: 3075: 3071: 3036: 3032: 2992: 2986: 2982: 2973: 2969: 2938: 2934: 2878: 2872: 2868: 2823: 2816: 2792:10.1.1.584.1430 2767: 2763: 2733: 2728: 2724: 2677: 2673: 2642: 2638: 2599: 2595: 2562: 2558: 2535: 2531: 2482: 2478: 2473: 2469: 2421: 2417: 2410: 2400:Springer-Verlag 2388: 2384: 2341: 2332: 2293: 2284: 2242: 2237: 2233: 2194: 2190: 2143: 2136: 2124: 2122: 2110: 2104: 2100: 2055: 2051: 2041: 2039: 2038:. 26 March 2018 2030: 2029: 2022: 2010: 2008: 1996: 1990: 1986: 1977: 1973: 1969: 1956: 1931:Biological pump 1927: 1904: 1896: 1880: 1863: 1830: 1806:eddy covariance 1759: 1752: 1748: 1744: 1713: 1709: 1651: 1610: 1603: 1602: 1601: 1541: 1533: 1532: 1531: 1496: 1486: 1485: 1484: 1437: 1427: 1426: 1425: 1414:Marine sediment 1398: 1388: 1387: 1386: 1347:Solubility pump 1335:Biological pump 1329: 1319: 1318: 1317: 1292: 1282: 1281: 1280: 1264:Carbon fixation 1249: 1234: 1224: 1223: 1222: 1203: 1187: 1140: 1138:Forms of carbon 1130: 1129: 1128: 1103: 1093: 1092: 1091: 1046: 1017: 988: 941: 888: 868: 810: 751: 709:of the Earth's 705:(caused by the 642: 619: 524:vascular plants 520: 498: 476:. These employ 400:vascular plants 396: 378: 376: 325: 270: 266: 243: 239: 235: 231: 227: 221: 217: 209: 205: 189: 181: 137:chemical energy 133: 24: 17: 12: 11: 5: 3437: 3427: 3426: 3421: 3416: 3401: 3400: 3340: 3291: 3226: 3203: 3200:on 2011-03-05. 3177:(6): 368–373. 3151: 3109: 3069: 3030: 3003:(2): 163–181. 2980: 2967: 2942:Marine Geology 2932: 2866: 2814: 2761: 2731: 2722: 2671: 2652:(3): 167–178. 2636: 2593: 2556: 2545:(2): 117–140. 2529: 2476: 2467: 2415: 2408: 2382: 2355:(8): 736–753. 2330: 2303:(2): 356–370. 2282: 2240: 2231: 2188: 2134: 2098: 2069:(3): 570–620. 2049: 2020: 1984: 1970: 1968: 1965: 1964: 1963: 1958: 1954: 1948: 1943: 1938: 1933: 1926: 1923: 1902: 1894: 1878: 1862: 1859: 1855:Neoproterozoic 1829: 1826: 1758: 1755: 1750: 1746: 1742: 1711: 1707: 1688: 1687: 1682: 1679: 1676: 1673: 1662: 1653: 1652: 1650: 1649: 1642: 1635: 1627: 1624: 1623: 1622: 1621: 1605: 1604: 1600: 1599: 1594: 1589: 1584: 1579: 1574: 1569: 1564: 1559: 1557:Deep biosphere 1554: 1549: 1543: 1542: 1539: 1538: 1535: 1534: 1530: 1529: 1527:Redfield ratio 1524: 1519: 1514: 1509: 1507:Nutrient cycle 1504: 1498: 1497: 1494:Biogeochemical 1492: 1491: 1488: 1487: 1483: 1482: 1477: 1472: 1467: 1466: 1465: 1460: 1450: 1448:Methanogenesis 1445: 1439: 1438: 1433: 1432: 1429: 1428: 1424: 1423: 1422: 1421: 1411: 1406: 1400: 1399: 1394: 1393: 1390: 1389: 1385: 1384: 1379: 1374: 1369: 1364: 1362:Microbial loop 1359: 1354: 1349: 1344: 1343: 1342: 1331: 1330: 1325: 1324: 1321: 1320: 1316: 1315: 1310: 1305: 1300: 1294: 1293: 1288: 1287: 1284: 1283: 1279: 1278: 1277: 1276: 1271: 1261: 1256: 1250: 1248: 1247: 1245:Chemosynthesis 1242: 1240:Photosynthesis 1236: 1235: 1230: 1229: 1226: 1225: 1221: 1220: 1215: 1210: 1204: 1202: 1201: 1200: 1199: 1188: 1186: 1185: 1179: 1173: 1167: 1161: 1155: 1149: 1142: 1141: 1136: 1135: 1132: 1131: 1127: 1126: 1121: 1116: 1111: 1105: 1104: 1101:Carbon dioxide 1099: 1098: 1095: 1094: 1090: 1089: 1084: 1079: 1074: 1069: 1064: 1059: 1054: 1048: 1047: 1044: 1043: 1040: 1039: 1031: 1030: 1024: 1023: 1016: 1013: 987: 984: 940: 937: 887: 884: 867: 864: 809: 806: 750: 747: 723:North Atlantic 703:seasonal cycle 696:critical depth 641: 638: 617: 516:Main article: 497: 494: 414:. Before the 375: 372: 360: 359: 358: 357: 356: 355: 324: 321: 268: 264: 250: 249: 248: 247: 246: 245: 241: 237: 233: 229: 225: 222: 219: 215: 207: 203: 192:chemosynthesis 187: 179: 176:carbon dioxide 132: 129: 80:chemosynthesis 76:photosynthesis 72:carbon dioxide 32:photoautotroph 15: 9: 6: 4: 3: 2: 3436: 3425: 3422: 3420: 3417: 3415: 3412: 3411: 3409: 3396: 3392: 3387: 3382: 3377: 3372: 3368: 3364: 3360: 3357: 3356: 3351: 3344: 3336: 3332: 3327: 3322: 3318: 3314: 3311:(1): GB1003. 3310: 3306: 3302: 3295: 3287: 3283: 3278: 3273: 3268: 3263: 3259: 3255: 3251: 3248: 3247: 3242: 3235: 3233: 3231: 3222: 3218: 3214: 3207: 3196: 3192: 3188: 3184: 3180: 3176: 3172: 3165: 3158: 3156: 3147: 3143: 3139: 3135: 3131: 3127: 3123: 3116: 3114: 3105: 3101: 3097: 3093: 3089: 3085: 3081: 3073: 3065: 3061: 3057: 3053: 3049: 3045: 3041: 3034: 3026: 3022: 3018: 3014: 3010: 3006: 3002: 2998: 2991: 2984: 2977: 2971: 2963: 2959: 2955: 2951: 2947: 2943: 2936: 2928: 2924: 2920: 2916: 2912: 2908: 2904: 2900: 2896: 2892: 2888: 2884: 2877: 2870: 2862: 2858: 2854: 2850: 2846: 2842: 2838: 2834: 2833: 2828: 2821: 2819: 2810: 2806: 2802: 2798: 2793: 2788: 2784: 2780: 2776: 2772: 2765: 2757: 2753: 2749: 2745: 2741: 2737: 2726: 2718: 2714: 2710: 2706: 2702: 2698: 2694: 2690: 2686: 2682: 2675: 2667: 2663: 2659: 2655: 2651: 2647: 2640: 2632: 2628: 2624: 2620: 2616: 2612: 2608: 2604: 2597: 2589: 2585: 2581: 2577: 2573: 2569: 2568: 2560: 2552: 2548: 2544: 2540: 2533: 2525: 2521: 2517: 2513: 2509: 2505: 2501: 2497: 2493: 2489: 2488: 2480: 2471: 2463: 2459: 2455: 2451: 2447: 2443: 2439: 2435: 2431: 2427: 2419: 2411: 2405: 2401: 2396: 2395: 2386: 2378: 2374: 2370: 2366: 2362: 2358: 2354: 2350: 2346: 2339: 2337: 2335: 2326: 2322: 2318: 2314: 2310: 2306: 2302: 2298: 2291: 2289: 2287: 2278: 2274: 2270: 2266: 2262: 2258: 2254: 2250: 2246: 2235: 2227: 2223: 2219: 2215: 2211: 2207: 2203: 2199: 2192: 2184: 2180: 2175: 2170: 2165: 2160: 2156: 2152: 2148: 2141: 2139: 2131: 2120: 2116: 2109: 2102: 2094: 2090: 2085: 2080: 2076: 2072: 2068: 2064: 2060: 2053: 2037: 2033: 2027: 2025: 2017: 2006: 2002: 1995: 1988: 1981: 1975: 1971: 1962: 1959: 1952: 1949: 1947: 1944: 1942: 1939: 1937: 1934: 1932: 1929: 1928: 1922: 1920: 1916: 1912: 1908: 1900: 1892: 1888: 1884: 1876: 1872: 1867: 1858: 1856: 1852: 1848: 1844: 1840: 1836: 1825: 1822: 1817: 1815: 1811: 1807: 1802: 1800: 1796: 1792: 1788: 1784: 1780: 1776: 1772: 1768: 1764: 1754: 1739: 1736: 1731: 1727: 1725: 1721: 1717: 1704: 1702: 1698: 1694: 1686: 1683: 1680: 1677: 1674: 1671: 1667: 1663: 1660: 1659: 1658: 1648: 1643: 1641: 1636: 1634: 1629: 1628: 1626: 1625: 1619: 1609: 1608: 1607: 1606: 1598: 1595: 1593: 1590: 1588: 1585: 1583: 1580: 1578: 1575: 1573: 1570: 1568: 1565: 1563: 1560: 1558: 1555: 1553: 1550: 1548: 1545: 1544: 1537: 1536: 1528: 1525: 1523: 1520: 1518: 1515: 1513: 1510: 1508: 1505: 1503: 1502:Marine cycles 1500: 1499: 1495: 1490: 1489: 1481: 1478: 1476: 1473: 1471: 1468: 1464: 1461: 1459: 1456: 1455: 1454: 1451: 1449: 1446: 1444: 1441: 1440: 1436: 1431: 1430: 1420: 1417: 1416: 1415: 1412: 1410: 1407: 1405: 1402: 1401: 1397: 1392: 1391: 1383: 1380: 1378: 1375: 1373: 1370: 1368: 1365: 1363: 1360: 1358: 1355: 1353: 1350: 1348: 1345: 1341: 1338: 1337: 1336: 1333: 1332: 1328: 1323: 1322: 1314: 1311: 1309: 1306: 1304: 1301: 1299: 1296: 1295: 1291: 1286: 1285: 1275: 1272: 1270: 1267: 1266: 1265: 1262: 1260: 1257: 1255: 1252: 1251: 1246: 1243: 1241: 1238: 1237: 1233: 1228: 1227: 1219: 1216: 1214: 1211: 1209: 1206: 1205: 1198: 1195: 1194: 1193: 1190: 1189: 1183: 1180: 1177: 1174: 1171: 1168: 1165: 1162: 1159: 1156: 1153: 1150: 1147: 1144: 1143: 1139: 1134: 1133: 1125: 1122: 1120: 1117: 1115: 1112: 1110: 1107: 1106: 1102: 1097: 1096: 1088: 1085: 1083: 1082:Boreal forest 1080: 1078: 1075: 1073: 1070: 1068: 1065: 1063: 1060: 1058: 1055: 1053: 1050: 1049: 1042: 1041: 1037: 1033: 1032: 1029: 1026: 1025: 1021: 1020: 1012: 1010: 1005: 1001: 992: 983: 981: 977: 973: 969: 961: 957: 953: 950: 945: 936: 934: 929: 925: 923: 919: 914: 910: 906: 897: 892: 883: 881: 877: 872: 863: 861: 857: 853: 852:North Pacific 849: 844: 842: 838: 834: 830: 826: 822: 818: 815: 814:micronutrient 805: 803: 799: 795: 791: 790:gravitational 787: 783: 779: 775: 767: 764: 760: 755: 746: 744: 740: 736: 732: 728: 724: 720: 716: 712: 708: 704: 699: 697: 693: 689: 685: 681: 676: 674: 670: 666: 662: 655: 651: 646: 637: 635: 631: 628: 624: 623:heat capacity 620: 613: 609: 600: 596: 594: 593: 588: 584: 580: 576: 572: 571:phytoplankton 568: 563: 561: 557: 553: 549: 545: 541: 537: 533: 529: 525: 519: 511: 507: 502: 493: 489: 487: 483: 479: 478:physiological 475: 471: 467: 463: 459: 456: 452: 448: 444: 443:transpiration 439: 437: 436:polar regions 433: 429: 425: 421: 417: 413: 409: 405: 401: 395: 371: 367: 365: 353: 352: 351: 350: 349: 348: 347: 345: 341: 337: 333: 329: 320: 318: 314: 310: 306: 305:heterotrophic 302: 298: 294: 293:nucleic acids 290: 286: 282: 278: 274: 262: 259: 255: 240:O + 4 S + 3 H 223: 213: 201: 200: 199: 198: 197: 196: 195: 193: 185: 177: 173: 165: 160: 156: 154: 150: 146: 142: 138: 128: 126: 122: 118: 114: 111: 107: 103: 99: 95: 91: 90: 85: 81: 77: 73: 69: 65: 61: 53: 49: 45: 41: 37: 33: 28: 22: 3358: 3353: 3343: 3308: 3304: 3294: 3249: 3244: 3221:10419/183215 3212: 3206: 3195:the original 3174: 3170: 3129: 3125: 3087: 3083: 3072: 3047: 3043: 3033: 3000: 2996: 2983: 2975: 2970: 2945: 2941: 2935: 2886: 2882: 2869: 2836: 2830: 2774: 2770: 2764: 2739: 2735: 2725: 2684: 2680: 2674: 2649: 2645: 2639: 2606: 2602: 2596: 2571: 2565: 2559: 2542: 2538: 2532: 2491: 2485: 2479: 2470: 2429: 2425: 2418: 2398:. New York: 2393: 2385: 2352: 2348: 2300: 2296: 2252: 2248: 2244: 2234: 2201: 2197: 2191: 2154: 2151:PLOS Biology 2150: 2129: 2123:. Retrieved 2118: 2114: 2101: 2066: 2062: 2052: 2040:. Retrieved 2035: 2015: 2009:. Retrieved 2004: 2000: 1987: 1979: 1974: 1946:Productivity 1915:biodiversity 1898: 1890: 1874: 1868: 1864: 1831: 1818: 1803: 1767:carbon cycle 1760: 1740: 1732: 1728: 1705: 1689: 1656: 1340:Martin curve 1327:Carbon pumps 1254:Calvin cycle 1208:Black carbon 1191: 1146:Total carbon 1087:Geochemistry 1028:Carbon cycle 997: 964: 930: 926: 901: 873: 869: 845: 841:aeolian dust 811: 782:silicic acid 771: 707:consequences 700: 677: 658: 605: 590: 564: 540:phototrophic 521: 490: 440: 397: 368: 361: 343: 327: 326: 261:carbohydrate 251: 211: 169: 164:Calvin cycle 149:lithotrophic 134: 120: 116: 87: 63: 57: 3424:Food chains 3090:: 200–225. 2777:(19): n/a. 1791:chlorophyll 1716:radioactive 1404:Carbon sink 1367:Viral shunt 1357:Marine snow 1213:Blue carbon 1067:Deep carbon 1062:Atmospheric 1052:Terrestrial 956:Flint Hills 886:Terrestrial 880:empirically 866:Measurement 802:thermocline 759:World Ocean 680:mixed layer 661:photic zone 650:kelp forest 614:rates (see 587:photic zone 552:brown algae 548:green algae 447:evaporation 428:temperature 299:to perform 108:, while in 3408:Categories 3171:BioScience 2736:Anal. Chem 2125:2015-06-01 2011:2015-06-01 1967:References 1887:irrigation 1875:actual NPP 1720:beta decay 1377:Whale pump 1372:Jelly pump 1352:Lipid pump 1077:Permafrost 1045:By regions 939:Grasslands 905:litterfall 903:turnover, 786:synthesise 743:hurricanes 711:axial tilt 692:dissipates 673:scattering 669:absorption 665:attenuated 654:macroalgae 560:seagrasses 544:eukaryotes 532:Eubacteria 512:microalgae 510:planktonic 482:anatomical 472:(CAM) and 412:liverworts 392:See also: 194:(bottom): 155:molecules. 98:food chain 94:autotrophs 3335:128460031 3146:0960-9822 3104:0009-2541 3064:1476-4687 3025:130057872 2927:206524740 2911:0036-8075 2787:CiteSeerX 2454:1354-1013 2317:1051-0761 2121:(6): 1–16 2007:(6): 1–16 1795:petagrams 1783:satellite 1763:biosphere 1693:stoppered 1666:carbon-14 1004:allometry 968:temperate 913:symbiotic 798:upwelling 778:phosphate 749:Nutrients 719:temperate 634:nutrients 630:insulates 612:metabolic 592:Sargassum 556:red algae 460:from the 458:nutrients 424:hydrology 416:evolution 364:ecosystem 275:or other 153:inorganic 141:organisms 42:Project, 36:autotroph 3395:17646656 3286:17616580 3215:: 1–18. 2919:20603496 2809:53465452 2756:19193192 2717:40425548 2709:17778634 2666:93847401 2631:10856212 2516:14826912 2462:55932276 2377:84344715 2325:37718097 2183:24143135 2093:10477309 2042:26 March 1925:See also 1907:cropland 1885:valley, 1871:land use 1837:, where 1775:habitats 1618:Category 972:tropical 970:) to 4 ( 848:Southern 821:cofactor 794:plankton 727:tropical 608:salinity 579:littoral 575:seaweeds 528:seaweeds 420:function 406:such as 313:food web 297:respired 295:, or be 281:proteins 145:sunlight 131:Overview 3419:Ecology 3386:1937509 3363:Bibcode 3313:Bibcode 3277:1911196 3254:Bibcode 3191:1310258 3005:Bibcode 2950:Bibcode 2891:Bibcode 2883:Science 2861:9657713 2841:Bibcode 2832:Science 2779:Bibcode 2689:Bibcode 2681:Science 2611:Bibcode 2603:Science 2576:Bibcode 2524:4299572 2496:Bibcode 2434:Bibcode 2357:Bibcode 2277:4325717 2257:Bibcode 2245:in situ 2226:4325562 2206:Bibcode 2174:3797030 1941:f-ratio 1911:pasture 1847:sulfate 1779:weather 1735:species 1463:Wetland 1435:Methane 1218:Kerogen 1119:Removal 1015:Aquatic 986:Forests 976:Wetland 954:in the 837:deserts 825:enzymes 774:nitrate 627:sea ice 569:called 536:archaea 506:diatoms 504:Marine 466:stomata 455:mineral 432:deserts 346:(NPP). 332:biomass 309:animals 273:glucose 258:reduced 254:polymer 60:ecology 40:SeaWiFS 3414:Plants 3393: 3383: 3333: 3284: 3274: 3189: 3144: 3102: 3062: 3044:Nature 3023: 2925: 2917: 2909: 2859: 2807: 2789: 2754: 2715: 2707: 2664: 2629: 2522: 2514: 2487:Nature 2460: 2452: 2406: 2375: 2323: 2315: 2275: 2249:Nature 2224: 2198:Nature 2181: 2171: 2091: 2084:103747 2081: 1909:; 22% 1839:barite 1835:barium 1757:Global 1616: 1597:CO2SYS 1458:Arctic 1197:marine 1057:Marine 994:Forest 980:Europe 960:Kansas 739:storms 735:basins 583:attach 451:leaves 408:mosses 291:, and 289:lipids 277:sugars 236:S → CH 182:) and 106:plants 52:GeoEye 3331:S2CID 3198:(PDF) 3187:JSTOR 3167:(PDF) 3021:S2CID 2993:(PDF) 2923:S2CID 2879:(PDF) 2805:S2CID 2713:S2CID 2662:S2CID 2520:S2CID 2458:S2CID 2373:S2CID 2321:S2CID 2273:S2CID 2222:S2CID 2111:(PDF) 1997:(PDF) 1893:(ΔNPP 1821:areal 1810:fAPAR 1718:(via 1592:C4MIP 1540:Other 1184:(PIC) 1178:(POC) 1172:(DIC) 1166:(DOC) 1160:(TIC) 1154:(TOC) 949:Konza 731:gyres 640:Light 317:Earth 263:, (CH 232:+ 4 H 218:O + O 212:light 184:water 121:gross 113:algae 100:. In 3391:PMID 3282:PMID 3142:ISSN 3100:ISSN 3060:ISSN 2915:PMID 2907:ISSN 2857:PMID 2752:PMID 2705:PMID 2627:PMID 2512:PMID 2450:ISSN 2404:ISBN 2313:ISSN 2179:PMID 2089:PMID 2044:2018 1901:(NPP 1883:Nile 1877:(NPP 1148:(TC) 1072:Soil 947:The 856:HNLC 850:and 831:and 817:iron 808:Iron 780:and 766:2009 688:wind 554:and 538:are 480:and 462:soil 426:and 410:and 301:work 214:→ CH 210:O + 162:The 84:life 50:and 44:NASA 3381:PMC 3371:doi 3359:104 3321:doi 3272:PMC 3262:doi 3250:104 3217:hdl 3179:doi 3134:doi 3092:doi 3088:513 3052:doi 3048:559 3013:doi 2958:doi 2946:184 2899:doi 2887:329 2849:doi 2837:281 2797:doi 2744:doi 2697:doi 2685:235 2654:doi 2619:doi 2607:288 2584:doi 2547:doi 2504:doi 2492:167 2442:doi 2365:doi 2305:doi 2265:doi 2253:383 2214:doi 2202:331 2169:PMC 2159:doi 2079:PMC 2071:doi 1879:act 1845:of 1819:In 1814:LAI 1812:or 1009:DBH 896:oak 894:An 823:in 741:or 686:by 682:. 671:or 434:or 256:of 228:+ O 206:+ H 178:(CO 119:or 117:net 92:or 58:In 3410:: 3389:. 3379:. 3369:. 3352:. 3329:. 3319:. 3309:22 3307:. 3303:. 3280:. 3270:. 3260:. 3243:. 3229:^ 3185:. 3175:36 3173:. 3169:. 3154:^ 3140:. 3130:33 3128:. 3124:. 3112:^ 3098:. 3086:. 3082:. 3058:. 3046:. 3042:. 3019:. 3011:. 2999:. 2995:. 2956:. 2944:. 2921:. 2913:. 2905:. 2897:. 2885:. 2881:. 2855:. 2847:. 2835:. 2829:. 2817:^ 2803:. 2795:. 2785:. 2775:32 2773:. 2750:. 2740:81 2738:. 2711:. 2703:. 2695:. 2683:. 2660:. 2650:10 2648:. 2625:. 2617:. 2605:. 2582:. 2572:32 2570:. 2543:18 2541:. 2518:. 2510:. 2502:. 2490:. 2456:. 2448:. 2440:. 2430:11 2428:. 2402:. 2371:. 2363:. 2351:. 2347:. 2333:^ 2319:. 2311:. 2301:11 2299:. 2285:^ 2271:. 2263:. 2251:. 2220:. 2212:. 2200:. 2177:. 2167:. 2155:11 2153:. 2149:. 2137:^ 2128:. 2117:. 2113:. 2087:. 2077:. 2067:63 2065:. 2061:. 2034:. 2023:^ 2014:. 2003:. 1999:. 1957:S) 1953:(H 1895:LC 1799:SI 1726:. 1710:CO 1274:C4 1269:C3 924:. 907:, 843:. 776:, 745:. 648:A 618:10 562:. 550:, 486:C3 474:C4 438:. 287:, 283:, 267:O) 224:CO 202:CO 186:(H 62:, 3397:. 3373:: 3365:: 3337:. 3323:: 3315:: 3288:. 3264:: 3256:: 3223:. 3219:: 3181:: 3148:. 3136:: 3106:. 3094:: 3066:. 3054:: 3027:. 3015:: 3007:: 3001:7 2964:. 2960:: 2952:: 2929:. 2901:: 2893:: 2863:. 2851:: 2843:: 2811:. 2799:: 2781:: 2758:. 2746:: 2732:2 2719:. 2699:: 2691:: 2668:. 2656:: 2633:. 2621:: 2613:: 2590:. 2586:: 2578:: 2553:. 2549:: 2526:. 2506:: 2498:: 2464:. 2444:: 2436:: 2412:. 2379:. 2367:: 2359:: 2353:8 2327:. 2307:: 2279:. 2267:: 2259:: 2241:2 2228:. 2216:: 2208:: 2185:. 2161:: 2119:3 2095:. 2073:: 2046:. 2005:3 1955:2 1903:0 1751:2 1747:2 1743:2 1712:3 1708:2 1646:e 1639:t 1632:v 1007:( 768:. 616:Q 269:n 265:2 244:O 242:2 238:2 234:2 230:2 226:2 220:2 216:2 208:2 204:2 188:2 180:2 54:. 46:/ 23:.

Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.