Primary production - Knowledge (XXG)

815:, 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. 1764:/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. 705:(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 503:

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.

512: 2027:"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. 1047: 765: 38: 610: 1624: 656: 882:

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.

170: 889:, 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 1877:

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.

1002: 902: 441:(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 1714:. 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. 955: 1932:,. 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. 1834:

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

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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

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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

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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

674:(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 398:

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;

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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.

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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.

647:, 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. 976:

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

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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,

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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

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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

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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

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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

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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).

1710:. 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 1843:

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

479:. 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 1701:

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

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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".

390: 985:) 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. 1017:

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

381:"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. 2485:

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

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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.

1756:/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 3049:

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).

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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

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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

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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".

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

869:(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 2780:

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".

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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".

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

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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

946:. During night, this technique measures all components of ecosystem respiration. This respiration is scaled to day-time values and further subtracted from NEE. 495:

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

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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.

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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

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

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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

709:. As long as there are adequate nutrients available, net primary production occurs whenever the mixed layer is shallower than the critical depth. 1741:

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

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

2418: 1597: 1924:) was devoted to human agriculture. This disproportionate amount reduces the energy available to other species, having a marked impact on 533:

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

2118: 1860:. Together these records suggest large shifts in primary production throughout Earth's past with notable rises associated with Earth's 1062: 404: 1725:) to infer primary production is most commonly used today because it is sensitive, and can be used in all ocean environments. As C is 392: 1707: 2612:

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

1706:. The first is analysed immediately and used to determine the initial oxygen concentration; usually this is done by performing a 865:

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

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

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

345:, that primary producers create in a given length of time. Some fraction of this fixed energy is used by primary producers for 717: 1852:

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

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

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

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

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

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

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

480: 1908:) of 9.6% across global land-mass. In addition to this, end consumption by people raises the total HANPP to 23.8% of 857:

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

569:, and a diverse group of unicellular groups. Vascular plants are also represented in the ocean by groups such as the 58: 807:, dead or fecal material), nutrients are constantly lost from the photic zone, and are only replenished by mixing or 643:

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

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

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

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

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

290:. These relatively simple molecules may be then used to further synthesise more complicated molecules, including 1587: 1207: 1019: 528: 3205: 2319: 1648: 537:

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

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

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

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

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

748:, light may only vary slightly across the year, and mixing may only occur episodically, such as during large 3252:"Quantifying and mapping the human appropriation of net primary production in Earth's terrestrial ecosystems" 1804:

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

93:, which uses the oxidation or reduction of inorganic chemical compounds as its source of energy. Almost all 1572: 1087: 831: 350: 1733:), it is relatively straightforward to measure its incorporation in organic material using devices such as 576:

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

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

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

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

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

311: 295: 112: 3090: 3434: 3373: 3323: 3264: 3015: 2960: 2901: 2851: 2789: 2699: 2621: 2586: 2506: 2444: 2367: 2267: 2216: 1504: 1406: 1162: 1067: 346: 307: 135: 2802: 2158:"Biotic and Human Vulnerability to Projected Changes in Ocean Biogeochemistry over the 21st Century" 3000: 1453: 1269: 1124: 686:

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

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

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

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

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

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

3401: 3312:"Farming the planet: 1. Geographic distribution of global agricultural lands in the year 2000" 3292: 3152: 3110: 3070: 2925: 2917: 2867: 2842: 2762: 2715: 2637: 2522: 2460: 2414: 2323: 2189: 2099: 1946: 1785: 1784:. However, quantifying primary production at this scale is difficult because of the range of 1726: 1463: 1242: 1077: 962: 851: 843: 839: 796: 773: 606:, but the vast majority of free-floating production takes place within microscopic organisms. 342: 120: 99: 42: 3429: 3391: 3381: 3331: 3282: 3272: 3227: 3189: 3144: 3102: 3062: 3023: 2968: 2909: 2859: 2807: 2754: 2707: 2664: 2629: 2594: 2557: 2514: 2497: 2452: 2375: 2315: 2275: 2224: 2179: 2169: 2089: 2081: 1961: 1429: 1323: 1318: 182: 78: 2887:"Terrestrial Gross Carbon Dioxide Uptake: Global Distribution and Covariation with Climate" 1046: 701:

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

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

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Differences in relative photosynthesis between plankton species under different irradiance

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

456:. The latter process (which is responsible for about 90% of water use) is driven by the 3377: 3327: 3268: 3019: 2964: 2905: 2855: 2793: 2703: 2625: 2590: 2510: 2448: 2371: 2271: 2220: 609: 3396: 3360: 3287: 3251: 2403: 2184: 2157: 1865: 1760:/Ar have the additional advantage of not needing incubations in closed containers and O 1703: 1567: 1537: 1517: 1458: 1372: 1255: 1250: 1111: 858: 706: 410: 202: 186: 90: 86: 82: 2972: 2456: 2094: 2069: 17: 3418: 3345: 3035: 2937: 2838:"Primary production of the Biosphere: Integrating Terrestrial and Oceanic Components" 2379: 1809: 1781: 1013:

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

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

2472: 2387: 2335: 655: 322:, then transfers these organic molecules (and the energy stored within them) up the 2534: 2287: 2236: 2085: 1925: 1777: 1350: 1264: 1218: 1156: 1038: 792: 776: 446: 271: 174: 3106: 2633: 2356:"Estimating net primary productivity from grassland biomass dynamics measurements" 3051:"Triple oxygen isotope evidence for limited mid-Proterozoic primary productivity" 2863: 2711: 2174: 1835:

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

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

1917: 1801: 1414: 1387: 1382: 1377: 1367: 1337: 1223: 1082: 966: 812: 769: 702: 690: 675: 671: 660: 597: 457: 438: 315: 2837: 2561: 353:"). The remaining fixed energy (i.e., mass of photosynthate) is referred to as 3148: 3066: 2598: 1916:). It is estimated that, in 2000, 34% of the Earth's ice-free land area (12% 1897: 1730: 1362: 915: 721: 694: 683: 664: 622: 562: 558: 550: 488: 159: 126:

predominate in this role. Ecologists distinguish primary production as either

108: 3156: 3114: 3074: 2921: 2464: 2327: 3386: 3277: 3132: 2913: 2668: 1793: 1773: 1676: 1014: 978: 931:, the total carbon dioxide produced by the ecosystem, can also be made with 923: 890: 808: 788: 753: 729: 644: 602: 554: 434: 426: 374: 163: 104: 46: 3405: 3296: 2929: 2766: 2719: 2641: 2526: 2193: 2103: 557:

are significant contributors to primary production in the ocean, including

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Synthesis of organic compounds from carbon dioxide by biological organisms

3091:"Claypool continued: Extending the isotopic record of sedimentary sulfate" 2871: 3336: 3311: 2811: 1881: 982: 804: 733: 618: 589: 570: 566: 553:, none are known to utilise oxygen-evolving photosynthesis. A number of 542: 538: 520: 468: 323: 155: 151: 3231: 3001:"Barium in Deep-Sea Sediment: A Geochemical Proxy for Paleoproductivity" 1880:

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

993:, annual mowing makes the annual biomass increment of wetlands evident. 3201: 1921: 1857: 1789: 1745: 1445: 1228: 986: 862: 800: 784: 737: 637: 585: 546: 492: 465: 291: 283: 264: 89:, which uses light as its source of energy, but it also occurs through 70: 50: 3224:

Proceedings of the International Conference: Ecology, Society, Economy

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

2758: 1001: 901: 341:(GPP) is the amount of chemical energy, typically expressed as carbon 2518: 2279: 2228: 1849: 1845: 1607: 990: 970: 847: 835: 713: 516: 442: 373:

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

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

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

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

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

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

846:. A major source of iron to the oceans is dust from the Earth's 1893: 1831: 866: 827: 741: 725: 698: 472: 461: 418: 94: 54: 2985:

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

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of plants. Transpiration allows plants to transport water and

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

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

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of seawater buffers temperature changes, and the formation of

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

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The technique of using C incorporation (added as labelled Na

632:), ranges less widely in the ocean than on land because the 584:. Larger autotrophs, such as the seagrasses and macroalgae ( 2745:/Ar measurements by equilibrator inlet mass spectrometry". 2320:

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

541:. They include photoautotrophs from a variety of groups. 989:

productivity (marshes and fens) is similarly measured. In

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but a minute fraction of primary production is driven by

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fluorescence kinetics (technique still a research topic)

134:, the former accounting for losses to processes such as 3175:"Human appropriation of the products of photosynthesis" 2068:

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

1864:(approximately 2.4 to 2.0 billion years ago) and the 724:), although wind magnitudes additionally have strong 1780:, estimating it at the global scale is important in 596:

to the underlying substrate but still be within the

2354:Scurlock, J.M.O.; Johnson, K.; Olson, R.J. (2002). 1808:

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

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Gross primary production and net primary production

2402: 621:can be). Similarly, temperature, while affecting 1815:Scaling ecosystem-level GPP estimations based on 592:zone and adjacent shallow waters, where they can 2999:Dymond, Jack; Suess, Erwin; Lyle, Mitch (1992). 1800:(NDVI) for terrestrial habitats and sea-surface 799:their cells and cellular machinery. Because of 85:. It principally occurs through the process of 2831: 2829: 1868:(approximately 1.0 to 0.54 billion years ago). 3133:"The geologic history of primary productivity" 413:, with a small fraction coming from algae and 3168: 3166: 2043:"Net Primary Productivity : Global Maps" 1649: 909:tree; a typical modern, terrestrial autotroph 8: 3245: 3243: 3241: 670:The sunlit zone of the ocean is called the 2119:"The Biological Productivity of the Ocean" 2037: 2035: 2005:"The Biological Productivity of the Ocean" 1928:, flows of carbon, water, and energy, and 1752:The methods based on stable isotopes and O 1656: 1642: 1029: 3395: 3385: 3335: 3286: 3276: 2801: 2183: 2173: 2093: 1892:). Although in some regions, such as the 1884:, results in various levels of impact on 850:, picked up and delivered by the wind as 803:sinking of particulate material (such as 185:from simpler inorganic compounds such as 2349: 2347: 2345: 768:Annual mean sea surface nitrate for the 499:plants (the majority of plant species). 388: 146:Primary production is the production of 36: 2301: 2299: 2297: 1982: 1788:on Earth, and because of the impact of 1593:Territorialisation of carbon governance 1037: 728:. Consequently, primary production in 433:of many factors, but principally local 162:organisms using the chemical energy of 2151: 2149: 1798:Normalized Difference Vegetation Index 1686:Stable isotopes of Oxygen (O, O and O) 314:. Consumption of primary producers by 2405:Primary Productivity of the Biosphere 1598:Total Carbon Column Observing Network 154:. The main source of this energy is 7: 2657:Limnology and Oceanography: Methods 2401:Leith, H.; Whittaker, R.H. (1975). 1692:Stable isotopes of Carbon (C and C) 922:, root exudates, and allocation to 795:are necessary for phytoplankton to 405:Terrestrial biological carbon cycle 263:In both cases, the end point is a 25: 2457:10.1111/j.1365-2486.2005.001002.x 2117:Sigman, D.M.; Hain, M.P. (2012). 2003:Sigman, D.M.; Hain, M.P. (2012). 1902:NPP reduction due to land changes 600:. There are exceptions, such as 3361:"Our share of the planetary pie" 2380:10.1046/j.1365-2486.2002.00512.x 1972:Lists of organisms by population 1623: 1622: 1045: 588:) are generally confined to the 1991:Terrestrial Global Productivity 1022:or BAI), and volume increment. 150:in organic compounds by living 41:Global oceanic and terrestrial 2156:Mora, C.; et al. (2013). 2086:10.1128/MMBR.63.3.570-620.1999 2070:"Bioenergetics of the Archaea" 1872:Human impact and appropriation 1558:Climate reconstruction proxies 893:determined conversion factor. 887:biochemically based techniques 838:involved in processes such as 282:, typically molecules such as 32:Primary production (economics) 1: 3366:Proc. Natl. Acad. Sci. U.S.A. 3257:Proc. Natl. Acad. Sci. U.S.A. 3107:10.1016/j.chemgeo.2019.02.030 2973:10.1016/S0025-3227(01)00286-9 2634:10.1126/science.288.5473.2028 1772:As primary production in the 678:down the water column by its 3316:Global Biogeochemical Cycles 2864:10.1126/science.281.5374.237 2712:10.1126/science.235.4785.199 2175:10.1371/journal.pbio.1001682 1776:is an important part of the 1528:Carbonate compensation depth 1193:Particulate inorganic carbon 481:Crassulacean acid metabolism 81:from atmospheric or aqueous 1749:them from their consumers. 1675:incorporation of inorganic 744:in the middle of the major 107:, and form the base of the 59:Goddard Space Flight Center 3451: 2578:Limnology and Oceanography 2126:Nature Education Knowledge 2012:Nature Education Knowledge 1839:Present and Past Estimates 1796:-derived estimates of the 1583:Carbon capture and storage 1187:Particulate organic carbon 1181:Dissolved inorganic carbon 920:volatile organic compounds 526: 402: 29: 3149:10.1016/j.cub.2023.09.040 3067:10.1038/s41586-018-0349-y 2599:10.4319/lo.1987.32.5.1085 2550:J. Cons. Int. Explor. Mer 2074:Microbiol. Mol. Biol. Rev 2047:earthobservatory.nasa.gov 1588:Carbon cycle re-balancing 944:eddy covariance technique 663:; an example of attached 529:Marine primary production 2989:, 13 (1958), pp. 153-212 2987:Geochim. Cosmochim. Acta 2562:10.1093/icesjms/18.2.117 2254:fugacity in response to 1563:Carbon-to-nitrogen ratio 1523:Carbonate–silicate cycle 1491:Carbon dioxide clathrate 1486:Clathrate gun hypothesis 1314:Net ecosystem production 1175:Dissolved organic carbon 1009:Methods used to measure 365:NPP = GPP - respiration 339:Gross primary production 3387:10.1073/pnas.0705190104 3278:10.1073/pnas.0704243104 2914:10.1126/science.1184984 2669:10.4319/lom.2012.10.167 2308:Ecological Applications 1993:, Academic Press, 33-59 1573:Deep Carbon Observatory 1033:Part of a series on the 351:maintenance respiration 1854:triple oxygen isotopes 1735:scintillation counters 1393:Continental shelf pump 1169:Total inorganic carbon 1135:Satellite measurements 1006: 973: 910: 830:. This is used as a 780: 667: 614: 524: 400: 385:Terrestrial production 355:net primary production 178: 113:terrestrial ecoregions 66: 18:Net primary production 2437:Global Change Biology 2360:Global Change Biology 2258:iron fertilization". 1862:Great Oxidation Event 1712:ecosystem respiration 1683:) into organic matter 1578:Global Carbon Project 1309:Ecosystem respiration 1004: 957: 933:gas flux measurements 929:ecosystem respiration 904: 767: 740:regions, such as the 658: 612: 578:microscopic organisms 514: 397: 326:, fueling all of the 296:complex carbohydrates 172: 40: 3337:10.1029/2007GB002952 3143:(21): 4741–4750.e5. 2812:10.1029/2005GL023459 1910:potential vegetation 1782:Earth system science 1407:Carbon sequestration 1163:Total organic carbon 732:regions such as the 347:cellular respiration 136:cellular respiration 77:is the synthesis of 30:For other uses, see 3378:2007PNAS..10412585F 3372:(31): 12585–12586. 3328:2008GBioC..22.1003R 3269:2007PNAS..10412942H 3263:(31): 12942–12947. 3020:1992PalOc...7..163D 2965:2002MGeol.184..189P 2906:2010Sci...329..834B 2856:1998Sci...281..237F 2794:2005GeoRL..3219605K 2704:1987Sci...235..199C 2626:2000Sci...288.2028L 2620:(5473): 2028–2031. 2591:1987LimOc..32.1085B 2511:1951Natur.167..684N 2449:2005GCBio..11.1424R 2372:2002GCBio...8..736S 2272:1996Natur.383..511C 2221:1988Natur.331..341M 1696:Oxygen/Argon Ratios 1454:Atmospheric methane 1420:Soil carbon storage 1270:Reverse Krebs cycle 1125:Ocean acidification 1011:forest productivity 415:non-vascular plants 330:'s living systems. 318:organisms, such as 115:, these are mainly 2782:Geophys. Res. Lett 1930:ecosystem services 1681:sodium bicarbonate 1679:(C in the form of 1533:Great Calcite Belt 1481:Aerobic production 1301:Carbon respiration 1243:Metabolic pathways 1203:Primary production 1007: 974: 911: 781: 726:spatial components 668: 615: 525: 507:Oceanic production 460:of water from the 401: 179: 138:, the latter not. 121:aquatic ecoregions 75:primary production 67: 3061:(7715): 613–616. 3028:10.1029/92PA00181 2900:(5993): 834–838. 2850:(5374): 237–240. 2759:10.1021/ac802300u 2698:(4785): 199–202. 2505:(4252): 684–685. 2420:978-0-387-07083-4 2266:(6600): 511–513. 2215:(6154): 341–343. 1947:Biomass (ecology) 1708:Winkler titration 1666: 1665: 1464:Methane emissions 1120:In the atmosphere 963:tallgrass prairie 844:nitrogen fixation 840:nitrate reduction 774:World Ocean Atlas 772:. Data from the 395: 183:organic molecules 177:of photosynthesis 100:primary producers 79:organic compounds 16:(Redirected from 3442: 3410: 3409: 3399: 3389: 3356: 3350: 3349: 3339: 3307: 3301: 3300: 3290: 3280: 3247: 3236: 3235: 3219: 3213: 3212: 3210: 3204:. Archived from 3179: 3170: 3161: 3160: 3128: 3119: 3118: 3095:Chemical Geology 3085: 3079: 3078: 3046: 3040: 3039: 3008:Paleoceanography 3005: 2996: 2990: 2983: 2977: 2976: 2959:(3–4): 189–206. 2948: 2942: 2941: 2891: 2882: 2876: 2875: 2833: 2824: 2823: 2805: 2777: 2771: 2770: 2753:(5): 1855–1864. 2738: 2732: 2731: 2687: 2681: 2680: 2652: 2646: 2645: 2609: 2603: 2602: 2585:(5): 1085–1098. 2572: 2566: 2565: 2545: 2539: 2538: 2519:10.1038/167684b0 2492: 2486: 2483: 2477: 2476: 2443:(9): 1424–1439. 2431: 2425: 2424: 2408: 2398: 2392: 2391: 2351: 2340: 2339: 2303: 2292: 2291: 2280:10.1038/383511a0 2247: 2241: 2240: 2229:10.1038/331341a0 2204: 2198: 2197: 2187: 2177: 2168:(10): e1001682. 2153: 2144: 2143: 2138: 2137: 2123: 2114: 2108: 2107: 2097: 2065: 2059: 2058: 2056: 2054: 2039: 2030: 2029: 2024: 2023: 2009: 2000: 1994: 1987: 1962:Hydrogen sulfide 1658: 1651: 1644: 1631: 1626: 1625: 1430:pelagic sediment 1324:Soil respiration 1319:Photorespiration 1049: 1030: 969:of northeastern 871:introducing iron 695:Turbulent mixing 519:; an example of 396: 21: 3450: 3449: 3445: 3444: 3443: 3441: 3440: 3439: 3415: 3414: 3413: 3358: 3357: 3353: 3309: 3308: 3304: 3249: 3248: 3239: 3221: 3220: 3216: 3208: 3194:10.2307/1310258 3177: 3172: 3171: 3164: 3137:Current Biology 3130: 3129: 3122: 3087: 3086: 3082: 3048: 3047: 3043: 3003: 2998: 2997: 2993: 2984: 2980: 2950: 2949: 2945: 2889: 2884: 2883: 2879: 2835: 2834: 2827: 2803:10.1.1.584.1430 2779: 2778: 2774: 2744: 2740: 2739: 2735: 2689: 2688: 2684: 2654: 2653: 2649: 2611: 2610: 2606: 2574: 2573: 2569: 2547: 2546: 2542: 2494: 2493: 2489: 2484: 2480: 2433: 2432: 2428: 2421: 2411:Springer-Verlag 2400: 2399: 2395: 2353: 2352: 2343: 2305: 2304: 2295: 2253: 2249: 2248: 2244: 2206: 2205: 2201: 2155: 2154: 2147: 2135: 2133: 2121: 2116: 2115: 2111: 2067: 2066: 2062: 2052: 2050: 2049:. 26 March 2018 2041: 2040: 2033: 2021: 2019: 2007: 2002: 2001: 1997: 1988: 1984: 1980: 1967: 1942:Biological pump 1938: 1915: 1907: 1891: 1874: 1841: 1817:eddy covariance 1770: 1763: 1759: 1755: 1724: 1720: 1662: 1621: 1614: 1613: 1612: 1552: 1544: 1543: 1542: 1507: 1497: 1496: 1495: 1448: 1438: 1437: 1436: 1425:Marine sediment 1409: 1399: 1398: 1397: 1358:Solubility pump 1346:Biological pump 1340: 1330: 1329: 1328: 1303: 1293: 1292: 1291: 1275:Carbon fixation 1260: 1245: 1235: 1234: 1233: 1214: 1198: 1151: 1149:Forms of carbon 1141: 1140: 1139: 1114: 1104: 1103: 1102: 1057: 1028: 999: 952: 899: 879: 821: 762: 720:of the Earth's 716:(caused by the 653: 630: 535:vascular plants 531: 509: 487:. These employ 411:vascular plants 407: 389: 387: 336: 281: 277: 254: 250: 246: 242: 238: 232: 228: 220: 216: 200: 192: 148:chemical energy 144: 35: 28: 23: 22: 15: 12: 11: 5: 3448: 3446: 3438: 3437: 3432: 3427: 3417: 3416: 3412: 3411: 3351: 3302: 3237: 3214: 3211:on 2011-03-05. 3188:(6): 368–373. 3162: 3120: 3080: 3041: 3014:(2): 163–181. 2991: 2978: 2953:Marine Geology 2943: 2877: 2825: 2772: 2742: 2733: 2682: 2663:(3): 167–178. 2647: 2604: 2567: 2556:(2): 117–140. 2540: 2487: 2478: 2426: 2419: 2393: 2366:(8): 736–753. 2341: 2314:(2): 356–370. 2293: 2251: 2242: 2199: 2145: 2109: 2080:(3): 570–620. 2060: 2031: 1995: 1981: 1979: 1976: 1975: 1974: 1969: 1965: 1959: 1954: 1949: 1944: 1937: 1934: 1913: 1905: 1889: 1873: 1870: 1866:Neoproterozoic 1840: 1837: 1769: 1766: 1761: 1757: 1753: 1722: 1718: 1699: 1698: 1693: 1690: 1687: 1684: 1673: 1664: 1663: 1661: 1660: 1653: 1646: 1638: 1635: 1634: 1633: 1632: 1616: 1615: 1611: 1610: 1605: 1600: 1595: 1590: 1585: 1580: 1575: 1570: 1568:Deep biosphere 1565: 1560: 1554: 1553: 1550: 1549: 1546: 1545: 1541: 1540: 1538:Redfield ratio 1535: 1530: 1525: 1520: 1518:Nutrient cycle 1515: 1509: 1508: 1505:Biogeochemical 1503: 1502: 1499: 1498: 1494: 1493: 1488: 1483: 1478: 1477: 1476: 1471: 1461: 1459:Methanogenesis 1456: 1450: 1449: 1444: 1443: 1440: 1439: 1435: 1434: 1433: 1432: 1422: 1417: 1411: 1410: 1405: 1404: 1401: 1400: 1396: 1395: 1390: 1385: 1380: 1375: 1373:Microbial loop 1370: 1365: 1360: 1355: 1354: 1353: 1342: 1341: 1336: 1335: 1332: 1331: 1327: 1326: 1321: 1316: 1311: 1305: 1304: 1299: 1298: 1295: 1294: 1290: 1289: 1288: 1287: 1282: 1272: 1267: 1261: 1259: 1258: 1256:Chemosynthesis 1253: 1251:Photosynthesis 1247: 1246: 1241: 1240: 1237: 1236: 1232: 1231: 1226: 1221: 1215: 1213: 1212: 1211: 1210: 1199: 1197: 1196: 1190: 1184: 1178: 1172: 1166: 1160: 1153: 1152: 1147: 1146: 1143: 1142: 1138: 1137: 1132: 1127: 1122: 1116: 1115: 1112:Carbon dioxide 1110: 1109: 1106: 1105: 1101: 1100: 1095: 1090: 1085: 1080: 1075: 1070: 1065: 1059: 1058: 1055: 1054: 1051: 1050: 1042: 1041: 1035: 1034: 1027: 1024: 998: 995: 951: 948: 898: 895: 878: 875: 820: 817: 761: 758: 734:North Atlantic 714:seasonal cycle 707:critical depth 652: 649: 628: 527:Main article: 508: 505: 425:. Before the 386: 383: 371: 370: 369: 368: 367: 366: 335: 332: 279: 275: 261: 260: 259: 258: 257: 256: 252: 248: 244: 240: 236: 233: 230: 226: 218: 214: 203:chemosynthesis 198: 190: 187:carbon dioxide 143: 140: 91:chemosynthesis 87:photosynthesis 83:carbon dioxide 43:photoautotroph 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 3447: 3436: 3433: 3431: 3428: 3426: 3423: 3422: 3420: 3407: 3403: 3398: 3393: 3388: 3383: 3379: 3375: 3371: 3368: 3367: 3362: 3355: 3352: 3347: 3343: 3338: 3333: 3329: 3325: 3322:(1): GB1003. 3321: 3317: 3313: 3306: 3303: 3298: 3294: 3289: 3284: 3279: 3274: 3270: 3266: 3262: 3259: 3258: 3253: 3246: 3244: 3242: 3238: 3233: 3229: 3225: 3218: 3215: 3207: 3203: 3199: 3195: 3191: 3187: 3183: 3176: 3169: 3167: 3163: 3158: 3154: 3150: 3146: 3142: 3138: 3134: 3127: 3125: 3121: 3116: 3112: 3108: 3104: 3100: 3096: 3092: 3084: 3081: 3076: 3072: 3068: 3064: 3060: 3056: 3052: 3045: 3042: 3037: 3033: 3029: 3025: 3021: 3017: 3013: 3009: 3002: 2995: 2992: 2988: 2982: 2979: 2974: 2970: 2966: 2962: 2958: 2954: 2947: 2944: 2939: 2935: 2931: 2927: 2923: 2919: 2915: 2911: 2907: 2903: 2899: 2895: 2888: 2881: 2878: 2873: 2869: 2865: 2861: 2857: 2853: 2849: 2845: 2844: 2839: 2832: 2830: 2826: 2821: 2817: 2813: 2809: 2804: 2799: 2795: 2791: 2787: 2783: 2776: 2773: 2768: 2764: 2760: 2756: 2752: 2748: 2737: 2734: 2729: 2725: 2721: 2717: 2713: 2709: 2705: 2701: 2697: 2693: 2686: 2683: 2678: 2674: 2670: 2666: 2662: 2658: 2651: 2648: 2643: 2639: 2635: 2631: 2627: 2623: 2619: 2615: 2608: 2605: 2600: 2596: 2592: 2588: 2584: 2580: 2579: 2571: 2568: 2563: 2559: 2555: 2551: 2544: 2541: 2536: 2532: 2528: 2524: 2520: 2516: 2512: 2508: 2504: 2500: 2499: 2491: 2488: 2482: 2479: 2474: 2470: 2466: 2462: 2458: 2454: 2450: 2446: 2442: 2438: 2430: 2427: 2422: 2416: 2412: 2407: 2406: 2397: 2394: 2389: 2385: 2381: 2377: 2373: 2369: 2365: 2361: 2357: 2350: 2348: 2346: 2342: 2337: 2333: 2329: 2325: 2321: 2317: 2313: 2309: 2302: 2300: 2298: 2294: 2289: 2285: 2281: 2277: 2273: 2269: 2265: 2261: 2257: 2246: 2243: 2238: 2234: 2230: 2226: 2222: 2218: 2214: 2210: 2203: 2200: 2195: 2191: 2186: 2181: 2176: 2171: 2167: 2163: 2159: 2152: 2150: 2146: 2142: 2131: 2127: 2120: 2113: 2110: 2105: 2101: 2096: 2091: 2087: 2083: 2079: 2075: 2071: 2064: 2061: 2048: 2044: 2038: 2036: 2032: 2028: 2017: 2013: 2006: 1999: 1996: 1992: 1986: 1983: 1977: 1973: 1970: 1963: 1960: 1958: 1955: 1953: 1950: 1948: 1945: 1943: 1940: 1939: 1935: 1933: 1931: 1927: 1923: 1919: 1911: 1903: 1899: 1895: 1887: 1883: 1878: 1871: 1869: 1867: 1863: 1859: 1855: 1851: 1847: 1838: 1836: 1833: 1828: 1826: 1822: 1818: 1813: 1811: 1807: 1803: 1799: 1795: 1791: 1787: 1783: 1779: 1775: 1767: 1765: 1750: 1747: 1742: 1738: 1736: 1732: 1728: 1715: 1713: 1709: 1705: 1697: 1694: 1691: 1688: 1685: 1682: 1678: 1674: 1671: 1670: 1669: 1659: 1654: 1652: 1647: 1645: 1640: 1639: 1637: 1636: 1630: 1620: 1619: 1618: 1617: 1609: 1606: 1604: 1601: 1599: 1596: 1594: 1591: 1589: 1586: 1584: 1581: 1579: 1576: 1574: 1571: 1569: 1566: 1564: 1561: 1559: 1556: 1555: 1548: 1547: 1539: 1536: 1534: 1531: 1529: 1526: 1524: 1521: 1519: 1516: 1514: 1513:Marine cycles 1511: 1510: 1506: 1501: 1500: 1492: 1489: 1487: 1484: 1482: 1479: 1475: 1472: 1470: 1467: 1466: 1465: 1462: 1460: 1457: 1455: 1452: 1451: 1447: 1442: 1441: 1431: 1428: 1427: 1426: 1423: 1421: 1418: 1416: 1413: 1412: 1408: 1403: 1402: 1394: 1391: 1389: 1386: 1384: 1381: 1379: 1376: 1374: 1371: 1369: 1366: 1364: 1361: 1359: 1356: 1352: 1349: 1348: 1347: 1344: 1343: 1339: 1334: 1333: 1325: 1322: 1320: 1317: 1315: 1312: 1310: 1307: 1306: 1302: 1297: 1296: 1286: 1283: 1281: 1278: 1277: 1276: 1273: 1271: 1268: 1266: 1263: 1262: 1257: 1254: 1252: 1249: 1248: 1244: 1239: 1238: 1230: 1227: 1225: 1222: 1220: 1217: 1216: 1209: 1206: 1205: 1204: 1201: 1200: 1194: 1191: 1188: 1185: 1182: 1179: 1176: 1173: 1170: 1167: 1164: 1161: 1158: 1155: 1154: 1150: 1145: 1144: 1136: 1133: 1131: 1128: 1126: 1123: 1121: 1118: 1117: 1113: 1108: 1107: 1099: 1096: 1094: 1093:Boreal forest 1091: 1089: 1086: 1084: 1081: 1079: 1076: 1074: 1071: 1069: 1066: 1064: 1061: 1060: 1053: 1052: 1048: 1044: 1043: 1040: 1036: 1032: 1031: 1025: 1023: 1021: 1016: 1012: 1003: 996: 994: 992: 988: 984: 980: 972: 968: 964: 961: 956: 949: 947: 945: 940: 936: 934: 930: 925: 921: 917: 908: 903: 896: 894: 892: 888: 883: 876: 874: 872: 868: 864: 863:North Pacific 860: 855: 853: 849: 845: 841: 837: 833: 829: 826: 825:micronutrient 818: 816: 814: 810: 806: 802: 801:gravitational 798: 794: 790: 786: 778: 775: 771: 766: 759: 757: 755: 751: 747: 743: 739: 735: 731: 727: 723: 719: 715: 710: 708: 704: 700: 696: 692: 687: 685: 681: 677: 673: 666: 662: 657: 650: 648: 646: 642: 639: 635: 634:heat capacity 631: 624: 620: 611: 607: 605: 604: 599: 595: 591: 587: 583: 582:phytoplankton 579: 574: 572: 568: 564: 560: 556: 552: 548: 544: 540: 536: 530: 522: 518: 513: 506: 504: 500: 498: 494: 490: 489:physiological 486: 482: 478: 474: 470: 467: 463: 459: 455: 454:transpiration 450: 448: 447:polar regions 444: 440: 436: 432: 428: 424: 420: 416: 412: 406: 384: 382: 378: 376: 364: 363: 362: 361: 360: 359: 358: 356: 352: 348: 344: 340: 333: 331: 329: 325: 321: 317: 316:heterotrophic 313: 309: 305: 304:nucleic acids 301: 297: 293: 289: 285: 273: 270: 266: 251:O + 4 S + 3 H 234: 224: 212: 211: 210: 209: 208: 207: 206: 204: 196: 188: 184: 176: 171: 167: 165: 161: 157: 153: 149: 141: 139: 137: 133: 129: 125: 122: 118: 114: 110: 106: 102: 101: 96: 92: 88: 84: 80: 76: 72: 64: 60: 56: 52: 48: 44: 39: 33: 19: 3369: 3364: 3354: 3319: 3315: 3305: 3260: 3255: 3232:10419/183215 3223: 3217: 3206:the original 3185: 3181: 3140: 3136: 3098: 3094: 3083: 3058: 3054: 3044: 3011: 3007: 2994: 2986: 2981: 2956: 2952: 2946: 2897: 2893: 2880: 2847: 2841: 2785: 2781: 2775: 2750: 2746: 2736: 2695: 2691: 2685: 2660: 2656: 2650: 2617: 2613: 2607: 2582: 2576: 2570: 2553: 2549: 2543: 2502: 2496: 2490: 2481: 2440: 2436: 2429: 2409:. 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