Photosynthetic efficiency

The photosynthetic efficiency is the fraction of light energy converted into chemical energy during photosynthesis in plants and algae. Photosynthesis can be described by the simplified chemical reaction

6H2O + 6CO2 + energy → C6H12O6 + 6O2

where C6H12O6 is glucose (which is subsequently transformed into other sugars, cellulose, lignin, and so forth). The value of the photosynthetic efficiency is dependent on how light energy is defined – it depends on whether we count only the light that is absorbed, and on what kind of light is used (see Photosynthetically active radiation). It takes eight (or perhaps 10 or more[1]) photons to utilize one molecule of CO2. The Gibbs free energy for converting a mole of CO2 to glucose is 114 kcal, whereas eight moles of photons of wavelength 600 nm contains 381 kcal, giving a nominal efficiency of 30%.[2] However, photosynthesis can occur with light up to wavelength 720 nm so long as there is also light at wavelengths below 680 nm to keep Photosystem II operating (see Chlorophyll). Using longer wavelengths means less light energy is needed for the same number of photons and therefore for the same amount of photosynthesis. For actual sunlight, where only 45% of the light is in the photosynthetically active wavelength range, the theoretical maximum efficiency of solar energy conversion is approximately 11%. In actuality, however, plants do not absorb all incoming sunlight (due to reflection, respiration requirements of photosynthesis and the need for optimal solar radiation levels) and do not convert all harvested energy into biomass, which results in a maximum overall photosynthetic efficiency of 3 to 6% of total solar radiation.[1] If photosynthesis is inefficient, excess light energy must be dissipated to avoid damaging the photosynthetic apparatus. Energy can be dissipated as heat (non-photochemical quenching), or emitted as chlorophyll fluorescence.

Typical efficiencies

Plants

Quoted values sunlight-to-biomass efficiency

Plant Efficiency
Plants, typical 0.1%[3]
0.2–2%[4]3.5-4.3%[5]
Typical crop plants 1–2%[3]

The following is a breakdown of the energetics of the photosynthesis process from Photosynthesis by Hall and Rao:[6]

Starting with the solar spectrum falling on a leaf,
47% lost due to photons outside the 400–700 nm active range (chlorophyll utilizes photons between 400 and 700 nm, extracting the energy of one 700 nm photon from each one)
30% of the in-band photons are lost due to incomplete absorption or photons hitting components other than chloroplasts
24% of the absorbed photon energy is lost due to degrading short wavelength photons to the 700 nm energy level
68% of the utilized energy is lost in conversion into d-glucose
35–45% of the glucose is consumed by the leaf in the processes of dark and photo respiration

Stated another way:
100% sunlight → non-bioavailable photons waste is 47%, leaving
53% (in the 400–700 nm range) → 30% of photons are lost due to incomplete absorption, leaving
37% (absorbed photon energy) → 24% is lost due to wavelength-mismatch degradation to 700 nm energy, leaving
28.2% (sunlight energy collected by chlorophyll) → 32% efficient conversion of ATP and NADPH to d-glucose, leaving
9% (collected as sugar) → 35–40% of sugar is recycled/consumed by the leaf in dark and photo-respiration, leaving
5.4% net leaf efficiency.

Many plants lose much of the remaining energy on growing roots. Most crop plants store ~0.25% to 0.5% of the sunlight in the product (corn kernels, potato starch, etc.).

Photosynthesis increases linearly with light intensity at low intensity, but at higher intensity this is no longer the case (see Photosynthesis-irradiance curve). Above about 10,000 lux or ~100 watts/square meter the rate no longer increases. Thus, most plants can only utilize ~10% of full mid-day sunlight intensity.[6] This dramatically reduces average achieved photosynthetic efficiency in fields compared to peak laboratory results. However, real plants (as opposed to laboratory test samples) have lots of redundant, randomly oriented leaves. This helps to keep the average illumination of each leaf well below the mid-day peak enabling the plant to achieve a result closer to the expected laboratory test results using limited illumination.

Only if the light intensity is above a plant specific value, called the compensation point the plant assimilates more carbon and releases more oxygen by photosynthesis than it consumes by cellular respiration for its own current energy demand.
Photosynthesis measurement systems are not designed to directly measure the amount of light absorbed by the leaf. Nevertheless, the light response curves that the class produces do allow comparisons in photosynthetic efficiency between plants.

Algae and other monocellular organisms

From a 2010 study by the University of Maryland, photosynthesizing Cyanobacteria have been shown to be a significant species in the global carbon cycle, accounting for 20–30% of Earth's photosynthetic productivity and convert solar energy into biomass-stored chemical energy at the rate of ~450 TW.[7]

Efficiencies of various biofuel crops

Popular choices for plant biofuels include: oil palm, soybean, castor oil, sunflower oil, safflower oil, corn ethanol, and sugar cane ethanol.

An analysis of a proposed Hawaiian oil palm plantation claimed to yield 600 gallons of biodiesel per acre per year. That comes to 2835 watts per acre or 0.7 W/m2.[8] Typical insolation in Hawaii is around 5.5 kWh/(m2day) or 230 W/m2.[9] For this particular oil palm plantation, if it delivered the claimed 600 gallons of biodiesel per acre per year, would be converting 0.3% of the incident solar energy to chemical fuel. Total photosynthetic efficiency would include more than just the biodiesel oil, so this 0.3% number is something of a lower bound.

Contrast this with a typical photovoltaic installation,[10] which would produce an average of roughly 22 W/m2 (roughly 10% of the average insolation), throughout the year. Furthermore, the photovoltaic panels would produce electricity, which is a high-quality form of energy, whereas converting the biodiesel into mechanical energy entails the loss of a large portion of the energy. On the other hand, a liquid fuel is much more convenient for a vehicle than electricity, which has to be stored in heavy, expensive batteries.

Most crop plants store ~0.25% to 0.5% of the sunlight in the product (corn kernels, potato starch, etc.), sugar cane is exceptional in several ways to yield peak storage efficiencies of ~8%.

Ethanol fuel in Brazil has a calculation that results in: "Per hectare per year, the biomass produced corresponds to 0.27 TJ. This is equivalent to 0.86 W/m2. Assuming an average insolation of 225 W/m2, the photosynthetic efficiency of sugar cane is 0.38%." Sucrose accounts for little more than 30% of the chemical energy stored in the mature plant; 35% is in the leaves and stem tips, which are left in the fields during harvest, and 35% are in the fibrous material (bagasse) left over from pressing.

C3 vs. C4 and CAM plants

C3 plants use the Calvin cycle to fix carbon. C4 plants use a modified Calvin cycle in which they separate Ribulose-1,5-bisphosphate carboxylase oxygenase (RuBisCO) from atmospheric oxygen, fixing carbon in their mesophyll cells and using oxaloacetate and malate to ferry the fixed carbon to RuBisCO and the rest of the Calvin cycle enzymes isolated in the bundle-sheath cells. The intermediate compounds both contain four carbon atoms, which gives C4. In Crassulacean acid metabolism (CAM), time isolates functioning RuBisCo (and the other Calvin cycle enzymes) from high oxygen concentrations produced by photosynthesis, in that O2 is evolved during the day, and allowed to dissipate then, while at night atmospheric CO2 is taken up and stored as malic or other acids. During the day, CAM plants close stomata and use stored acids as carbon sources for sugar, etc. production.

The C3 pathway requires 18 ATP and 12 NADPH for the synthesis of one molecule of glucose (3 ATP + 2 NADPH per CO2 fixed) while the C4 pathway requires 30 ATP and 12 NADPH (C3 + 2 ATP per CO2 fixed). In addition, we can take into account that each NADPH is equivalent to 3 ATP, that means both pathways require 36 additional (equivalent of) ATP[11] [better citation needed]. Despite this reduced ATP efficiency, C4 is an evolutionary advancement, adapted to areas of high levels of light, where the reduced ATP efficiency is more than offset by the use of increased light. The ability to thrive despite restricted water availability maximizes the ability to use available light. The simpler C3 cycle which operates in most plants is adapted to wetter darker environments, such as many northern latitudes.. Corn, sugar cane, and sorghum are C4 plants. These plants are economically important in part because of their relatively high photosynthetic efficiencies compared to many other crops. Pineapple is a CAM plant.

Research

Photorespiration

One efficiency-focused research topic is improving the efficiency of photorespiration. Around 25 percent of the time RuBisCO incorrectly collects oxygen molecules instead of CO
2
, creating CO
2
and ammonia that disrupt the photosynthesis process. Plants remove these byproducts via photorespiration, requiring energy and nutrients that would otherwise increase photosynthetic output. In C3 plants photorespiration can consume 20-50% of photosynthetic energy. The research shortened photosynthetic pathways in tobacco. Engineered crops grew taller and faster, yielding up to 40 percent more biomass. The study employed synthetic biology to construct new metabolic pathways and assessed their efficiency with and without transporter RNAi. The most efficient pathway increased light-use efficiency by 17%.[12]

See also

References

  1. ^ a b Renewable biological systems for unsustainable energy production. FAO Agricultural Services Bulletins (1997).
  2. ^ Stryer, Lubert (1981). Biochemistry (2nd ed.). p. 448. ISBN 978-0-7167-1226-8.
  3. ^ a b Govindjee, What is photosynthesis?
  4. ^ The Green Solar Collector; converting sunlight into algal biomass Wageningen University project (2005—2008)
  5. ^ Blankenship, Robert E.; Tiede, David M.; Barber, James; Brudvig, Gary W.; Fleming, Graham; Ghirardi, Maria; Gunner, M. R.; Junge, Wolfgang; Kramer, David M. (2011-05-13). "Comparing Photosynthetic and Photovoltaic Efficiencies and Recognizing the Potential for Improvement". Science. 332 (6031): 805–809. doi:10.1126/science.1200165. ISSN 0036-8075. PMID 21566184.
  6. ^ a b David Oakley Hall; K. K. Rao; Institute of Biology (1999). Photosynthesis. Cambridge University Press. ISBN 978-0-521-64497-6. Retrieved 3 November 2011.
  7. ^ Pisciotta JM, Zou Y, Baskakov IV (2010). "Light-Dependent Electrogenic Activity of Cyanobacteria". PLoS ONE. 5 (5): e10821. doi:10.1371/journal.pone.0010821. PMC 2876029. PMID 20520829.
  8. ^ Biodiesel Fuel. Ag.ndsu.edu.. Retrieved on 2011-11-03.
  9. ^ PVWATTS: Hawaii. Rredc.nrel.gov. Retrieved on 2011-11-03.
  10. ^ NREL: In My Backyard (IMBY) Home Page. Nrel.gov (2010-12-23). Retrieved on 2011-11-03.
  11. ^ "Biology –C4 Cycle - askIITians". www.askiitians.com.
  12. ^ South, Paul F.; Cavanagh, Amanda P.; Liu, Helen W.; Ort, Donald R. (2019-01-04). "Synthetic glycolate metabolism pathways stimulate crop growth and productivity in the field". Science. 363 (6422): eaat9077. doi:10.1126/science.aat9077. ISSN 0036-8075.
Cascade effect (ecology)

An ecological cascade effect is a series of secondary extinctions that is triggered by the primary extinction of a key species in an ecosystem. Secondary extinctions are likely to occur when the threatened species are: dependent on a few specific food sources, mutualistic (dependent on the key species in some way), or forced to coexist with an invasive species that is introduced to the ecosystem. Species introductions to a foreign ecosystem can often devastate entire communities, and even entire ecosystems. These exotic species monopolize the ecosystem's resources, and since they have no natural predators to decrease their growth, they are able to increase indefinitely. Olsen et al. showed that exotic species have caused lake and estuary ecosystems to go through cascade effects due to loss of algae, crayfish, mollusks, fish, amphibians, and birds. However, the principal cause of cascade effects is the loss of top predators as the key species. As a result of this loss, a dramatic increase (ecological release) of prey species occurs. The prey is then able to overexploit its own food resources, until the population numbers decrease in abundance, which can lead to extinction. When the prey's food resources disappear, they starve and may go extinct as well. If the prey species is herbivorous, then their initial release and exploitation of the plants may result in a loss of plant biodiversity in the area. If other organisms in the ecosystem also depend upon these plants as food resources, then these species may go extinct as well. An example of the cascade effect caused by the loss of a top predator is apparent in tropical forests. When hunters cause local extinctions of top predators, the predators' prey's population numbers increase, causing an overexploitation of a food resource and a cascade effect of species loss. Recent studies have been performed on approaches to mitigate extinction cascades in food-web networks.

Chlorella

Chlorella is a genus of single-celled green algae belonging to the division Chlorophyta. It is spherical in shape, about 2 to 10 μm in diameter, and is without flagella. Chlorella contains the green photosynthetic pigments chlorophyll-a and -b in its chloroplast. Through photosynthesis, it multiplies rapidly, requiring only carbon dioxide, water, sunlight, and a small amount of minerals to reproduce.The name Chlorella is taken from the Greek χλώρος, chloros, meaning green, and the Latin diminutive suffix ella, meaning small. German biochemist and cell physiologist Otto Heinrich Warburg, awarded with the Nobel Prize in Physiology or Medicine in 1931 for his research on cell respiration, also studied photosynthesis in Chlorella. In 1961, Melvin Calvin of the University of California received the Nobel Prize in Chemistry for his research on the pathways of carbon dioxide assimilation in plants using Chlorella.

Many people believe Chlorella can serve as a potential source of food and energy because its photosynthetic efficiency can, in theory, reach 8%, which exceeds that of other highly efficient crops such as sugar cane.

Crassulacean acid metabolism

Crassulacean acid metabolism, also known as CAM photosynthesis, is a carbon fixation pathway that evolved in some plants as an adaptation to arid conditions. In a plant using full CAM, the stomata in the leaves remain shut during the day to reduce evapotranspiration, but open at night to collect and allow carbon dioxide (CO2) to diffuse into the mesophyll cells. The CO2 is stored as the four-carbon acid malate in vacuoles at night, and then in the daytime, the malate is transported to chloroplasts where it is converted back to CO2, which is then used during photosynthesis. The pre-collected CO2 is concentrated around the enzyme RuBisCO, increasing photosynthetic efficiency. The mechanism was first discovered in plants of the family Crassulaceae.

Ecological thinning

Ecological thinning is a silvicultural technique used in forest management that involves cutting trees to improve functions of a forest other than timber production.

Although thinning originated as a man-made forest management tool, aimed at increasing timber yields, the shift from production forests to multifunctional forests brought with it the cutting of trees to manipulate an ecosystem for various reasons, ranging from removing non-native species from a plot to removing poplars growing on a riverside beach aimed at recreational use.

Since the 1970s, leaving the thinned trees on the forest floor has become an increasingly common policy: wood can be decomposed in a more natural fashion, playing an important role in increasing biodiversity by providing habitat to various invertebrates, birds and small mammals. Many fungi (e.g. Calocera viscosa) and mosses are saproxylic or epixylic as well (e.g. Marchantiophyta) – some moss species completing their entire life-cycle on a single log.

Where trees are managed under a commercial regime, competition is reduced by removing adjacent stems that exhibit less favourable timber quality potential. When left in a natural state trees will "self-thin", but this process can be unreliable in some circumstances. Examples of this can be found in the Buxus – Ironbark forests and woodlands of Victoria (Australia) where a large proportion of trees are coppice, resultant from timber cutting in decades gone by.

FLEX (satellite)

The FLuorescence EXplorer (FLEX) is a planned mission by the European Space Agency to launch a satellite to monitor the global steady-state chlorophyll fluorescence in terrestrial vegetation. FLEX was selected for funding on 19 November 2015 and will be launched on a Vega C rocket from Guiana Space Centre in 2023.

John Gamon

John A. Gamon is an American scientist currently working in Canada. His work using terrestrial vegetation spectral signatures to discern plant productivity and biodiversity has had a significant impact in the discipline of remote sensing, having published 95 papers and receiving 7,613 citations as of 2017. Gamon pioneered the use of the relationship between leaf xanthophyll cycle pigment content and spectral reflectance to improve satellite monitoring of photosynthesis. Gamon's seminal work resulted in the development of the Photochemical Reflectance Index (PRI). He trained under Nobel Prize laureate Christopher Field.

List of C4 plants

C4 plants use the C4 carbon fixation pathway to increase their photosynthetic efficiency by reducing or suppressing photorespiration, which mainly occurs under low atmospheric CO2 concentration, high light, high temperature, drought, and salinity. There are roughly 8,100 known C4 species, which belong to at least 61 distinct evolutionary lineages in 19 families (as per APG IV classification) of flowering plants. Among these are important crops such as maize, sorghum and sugarcane, but also weeds and invasive plants. Although only 3% of flowering plant species use C4 carbon fixation, they account for 23% of global primary production. The repeated, convergent C4 evolution from C3 ancestors has spurred hopes to bio-engineer the C4 pathway into C3 crops such as rice.C4 photosynthesis probably first evolved 30–35 million years ago in the Oligocene, and further origins occurred since, most of them in the last 15 million years. C4 plants are mainly found in tropical and warm-temperate regions, predominantly in open grasslands where they are often dominant. While most are graminoids, other growth forms such as forbs, vines, shrubs, and even some trees and aquatic plants are also known among C4 plants.C4 plants are usually identified by their higher 13C/12C isotopic ratio compared to C3 plants or their typical leaf anatomy. The distribution of C4 lineages among plants has been determined through phylogenetics and was considered well known as of 2016. Monocots – mainly grasses (Poaceae) and sedges (Cyperaceae) – account for around 80% of C4 species, but they are also found in the eudicots.The following list presents known C4 lineages by family, based on the overview by Sage (2016). They correspond to single species or clades thought to have acquired the C4 pathway independently. In some lineages that also include C3 and C3–C4 intermediate species, the C4 pathway may have evolved more than once.

Mango mealybug

Mango mealybug (Drosicha mangiferae), is a pest of mango crops in Asia. The nymphs and females suck plant sap from inflorescences, tender leaves, shoots and fruit peduncles. As a result, the infested inflorescences dry up, affects the fruit set, causing fruit drop. These bugs also exude honey dew over the mango tree leaves, on which sooty mold fungus develops reducing the photosynthetic efficiency of the tree. It is a polyphagous pest and is found on over 60 other plant species

Mesopredator release hypothesis

The mesopredator release hypothesis is an ecological theory used to describe the interrelated population dynamics between apex predators and mesopredators within an ecosystem, such that a collapsing population of the former results in dramatically-increased populations of the latter. This hypothesis describes the phenomenon of trophic cascade in specific terrestrial communities.

A mesopredator is a medium-sized, middle trophic level predator, which both preys and is preyed upon. Examples are raccoons, skunks, snakes, cownose rays, and small sharks.

Nanophytoplankton

Nanophytoplankton are particularly small phytoplankton with sizes between 2 and 20 µm. They are the autotrophic part of nanoplankton. Like other phytoplankton, nanophytoplankton are microscopic organisms that obtain energy through the process of photosynthesis and must therefore live in the upper sunlit layer of ocean or other bodies of water. These microscopic free-floating organisms, including algae, and cyanobacteria, fix large amounts of carbon which would otherwise be released as carbon dioxide.. The term nanophytoplankton is derived from the far more widely used term nannoplankton/nanoplankton.

Nepenthes talangensis

Nepenthes talangensis is a tropical pitcher plant endemic to Sumatra, where it grows in upper montane forest at elevations of 1800–2500 m above sea level.The specific epithet talangensis is derived from the name of Mount Talang, to which it is endemic, and the Latin ending -ensis, meaning "from".

Photochemical Reflectance Index

The Photochemical Reflectance Index (PRI) is a reflectance measurement developed by John Gamon during his tenure as a postdoctorate fellow supervised by Christopher Field at the Carnegie Institution for Science at Stanford University. The PRI is sensitive to changes in carotenoid pigments (e.g. xanthophyll pigments) in live foliage. Carotenoid pigments are indicative of photosynthetic light use efficiency, or the rate of carbon dioxide uptake by foliage per unit energy absorbed. As such, it is used in studies of vegetation productivity and stress. Because the PRI measures plant responses to stress, it can be used to assess general ecosystem health using satellite data or other forms of remote sensing. Applications include vegetation health in evergreen shrublands, forests, and agricultural crops prior to senescence. PRI is defined by the following equation using reflectance (ρ) at 531 and 570 nm wavelength:

Some authors use

The values range from –1 to 1.

Photosynthesis

Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that can later be released to fuel the organisms' activities. This chemical energy is stored in carbohydrate molecules, such as sugars, which are synthesized from carbon dioxide and water – hence the name photosynthesis, from the Greek φῶς, phōs, "light", and σύνθεσις, synthesis, "putting together". In most cases, oxygen is also released as a waste product. Most plants, most algae, and cyanobacteria perform photosynthesis; such organisms are called photoautotrophs. Photosynthesis is largely responsible for producing and maintaining the oxygen content of the Earth's atmosphere, and supplies all of the organic compounds and most of the energy necessary for life on Earth.Although photosynthesis is performed differently by different species, the process always begins when energy from light is absorbed by proteins called reaction centres that contain green chlorophyll pigments. In plants, these proteins are held inside organelles called chloroplasts, which are most abundant in leaf cells, while in bacteria they are embedded in the plasma membrane. In these light-dependent reactions, some energy is used to strip electrons from suitable substances, such as water, producing oxygen gas. The hydrogen freed by the splitting of water is used in the creation of two further compounds that serve as short-term stores of energy, enabling its transfer to drive other reactions: these compounds are reduced nicotinamide adenine dinucleotide phosphate (NADPH) and adenosine triphosphate (ATP), the "energy currency" of cells.

In plants, algae and cyanobacteria, long-term energy storage in the form of sugars is produced by a subsequent sequence of light-independent reactions called the Calvin cycle; some bacteria use different mechanisms, such as the reverse Krebs cycle, to achieve the same end. In the Calvin cycle, atmospheric carbon dioxide is incorporated into already existing organic carbon compounds, such as ribulose bisphosphate (RuBP). Using the ATP and NADPH produced by the light-dependent reactions, the resulting compounds are then reduced and removed to form further carbohydrates, such as glucose.

The first photosynthetic organisms probably evolved early in the evolutionary history of life and most likely used reducing agents such as hydrogen or hydrogen sulfide, rather than water, as sources of electrons. Cyanobacteria appeared later; the excess oxygen they produced contributed directly to the oxygenation of the Earth, which rendered the evolution of complex life possible. Today, the average rate of energy capture by photosynthesis globally is approximately 130 terawatts, which is about eight times the current power consumption of human civilization.

Photosynthetic organisms also convert around 100–115 billion tonnes (91-104 petagrams) of carbon into biomass per year.

Photosynthetic capacity

Photosynthetic capacity (Amax) is a measure of the maximum rate at which leaves are able to fix carbon during photosynthesis. It is typically measured as the amount of carbon dioxide that is fixed per metre squared per second, for example as μmol m−2 sec−1.

Productivity (ecology)

In ecology, productivity refers to the rate of generation of biomass in an ecosystem. It is usually expressed in units of mass per unit surface (or volume) per unit time, for instance grams per square metre per day (g m−2 d−1). The mass unit may relate to dry matter or to the mass of carbon generated. Productivity of autotrophs such as plants is called primary productivity, while that of heterotrophs such as animals is called secondary productivity.

Realizing Increased Photosynthetic Efficiency

Realizing Increased Photosynthetic Efficiency (RIPE) is a translational research project that is genetically engineering plants to photosynthesize more efficiently to increase crop yields. RIPE aims to increase agricultural production worldwide, particularly to help reduce hunger and poverty in Sub-Saharan Africa and Southeast Asia by sustainably improving the yield of key food crops including soybeans, rice, cassava and cowpeas. The RIPE project began in 2012, funded by a five-year, $25-million dollar grant from the Bill and Melinda Gates Foundation. In 2017, the project received a $45 million-dollar reinvestment from the Gates Foundation, Foundation for Food and Agriculture Research, and the UK Government's Department for International Development. In 2018, the Gates Foundation contributed an additional $13 million to accelerate the project's progress.

Ripe

Ripe or RIPE may refer to:

Realizing Increased Photosynthetic Efficiency (RIPE), a genetic engineering research project

Ripening, especially of fruit

Ripeness in viticulture, how the term "ripe" is used in viticulture and winemaking

RIPE, Réseaux IP Européens

RIPE NCC, the Regional Internet Registry (RIR) for Europe

RIPEMD, a family of cryptographic hash functions

Ripeness, a term in law

Ripe, East Sussex, in England

Ripe, Marche, in Italy

Ripe (Slug album), 2015 rock music album

Ripe (film), a 1996 drama released in 1997 by Mo Ogrodnik

Ripe (album), a 2007 album by Ben Lee

Wilhelm Ripe (1818–1885), German painter

Thalassiosira weissflogii

Thalassiosira weissflogii is a species of centric diatoms, a unicellular microalga. It is found in marine environments and also in inland waters in many parts of the world. It is actively studied because it may use C4-plant style strategies to increase its photosynthetic efficiency.

Zooxanthellae

Zooxanthellae are single-celled dinoflagellates that are able to live in symbiosis with diverse marine invertebrates including corals, jellyfish, and nudibranchs. Most known zooxanthellae are in the genus Symbiodinium, but some are known from the genus Amphidinium, and other taxa, as yet unidentified, may have similar endosymbiont affinities. Another group of unicellular eukaryotes that partake in similar endosymbiotic relationships in both marine and freshwater habitats are green algae zoochlorellae.Zooxanthellae are photosynthetic organisms, which contain chlorophyll a and chlorophyll c, as well as the dinoflagellate pigments peridinin and diadinoxanthin. These provide the yellowish and brownish colours typical of many of the host species. During the day, they provide their host with the organic carbon products of photosynthesis, sometimes providing up to 90% of their host's energy needs for metabolism, growth and reproduction. In return, they receive nutrients, carbon dioxide, and an elevated position with access to sunshine.

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