According to the Table, What Is the Optimum Temperature for the Production of Glucose in Plants?

Terrestrial plants are regularly subjected to strong temperature variations. These variations can achieve an amplitude of 40°C or even more, both in polar regions and in hot desert areas. Being rooted, they have reduced mobility and must cope with changes in their environment. The absorption of CO₂ by plants via photosynthesis is the gateway to carbon in the biosphere. What is the thermal amplitude that allows it to office? How does photosynthesis reacts to rapid and slow temperature variations? What is the multifariousness of responses? What are the physiological processes that limit it? Crucial questions tare o be considered in the context of global warming.

1. Plant product and climate alter

The current increment in greenhouse gas emissions will cause an increase in atmospheric temperature of 2 to 3°C in the adjacent 50 years (see A carbon bike disrupted by man activities). At the same time, heat waves and extreme estrus periods will exist more frequent and of longer duration [1]. Agricultural output and the functioning of forests will therefore exist greatly affected. Models based on big-scale observations indicate that, in the absence of agronomic adaptation, the decrease in ingather yields can reach 17% for each 1°C increase in the temperature of the growing flavour [2].

The production of higher plants depends in item (only not only [3]) on leaf photosynthesis (come across Shedding light on photosynthesis & The The path of carbon in photosynthesis). CO_ii enters the foliage where its reduction in the chloroplasts is accompanied past O_ii production. Its entry is near exclusively through the stomata (Effigy 1). For each molecule of CO_two absorbed, fifty to 300 molecules of water are transpired from the leaves, depending on the establish. This water allows, among other things, the cooling of the leafage (run across Focus Leafage transpiration and heat protection).

Figure one. During photosynthesis, CO2 is absorbed and O2 is released mainly through the stomatal opening (ostiole). The h2o vapour (transpiration of the leaf) passes mainly through the ostiole but also through the epidermis. Transpiration allows the leaf to cool down in the light. [Source: Writer'southward diagram]

The leaf is a converter of solar energy into chemical free energy and, like whatsoever energy converter, requires a permanent cooling system.

The climate changes that are currently occurring make information technology necessary to understand the effects of temperature on photosynthesis.

ii. The thermal optimum of photosynthesis

two.ane. Diagram of the thermal response

Photosynthetic CO₂ uptake varies with temperature. In most cases its response to temperature is rapidly reversible between most 10 and 34°C. In this range of temperatures it presents a maximum value: a thermal optimum.

Figure 2. Diagram of the variation of CO2 assimilation by an intact leaf. It highlights the temperature range in which  variations are generally rapidly reversible. [Source: Author's diagram]

Below ten°C and above 34°C plants start to set up up protective mechanisms. For these extreme values, CO_ii absorption is ofttimes unstable and can exist cancelled more or less quickly: the leaf is and then under stress (Figure two).

two.2. A thermal optimum based on the average temperature of the surround

Plants in common cold environments or with a cold growing flavor have a higher photosynthesis at low temperatures. Plants in warm environments, or growing during the warm season, accept a higher photosynthesis at high temperatures.

Figure 3. Deschampsia antarctica is ane of two flowering plants found in Antarctica. It is often subjected to negative temperatures. The snow that frequently covers it protects it from extreme temperatures. [Source: Lomvi2, CC By-SA 3.0, via Wikimedia Commons]

For example, the thermal optimum for CO₂ assimilation [4] in Deschampsia antarctica (Effigy 3) and Colobanthus quitensis, the only 2 Antarctic flowering plants, is between eight and 15°C, while it is around 45°C in Tridestomia oblongifolia, a warm desert plant from Key America. The latter species probably holds the world record for flowering plants in this respect.

two.3. Acclimatization to the thermal conditions of the environment

Figure 4. Variations in CO2 absorption as a office of foliage temperature, in a institute grown at 10°C (red) or 25°C. Measurements made on Pea, under a light close to saturation. CO2 content in ambient air: 390 ppm. [Source: Author's diagram]

Differences in the thermal response of photosynthesis are as well constitute in individuals of the same species growing at different temperatures. Figure 4 shows CO₂ assimilation in pea grown at ten or 25°C.

In the first case (cultivation at 10°C) the thermal optimum is about xvi°C, while it is higher than 25°C in the second (cultivation at 25°C). At low temperatures, CO₂ absorption is higher in plants grown at x°C.

In this case the adjustment to cool conditions is a gain for the establish.

ii.iv. Acclimatization tin be rapid

Figure 5. Remth (Hammada scoparia), a feature plant of the Wadi Rum desert (Jordan). [Source: Ji-Elle, CC By-SA iv.0, via Wikimedia Eatables]

For example, the photosynthesis of Hammada scoparia, a bush-league in the deserts of the Center Eastward (Negev, Wadi Rum) follows the seasonal variations in temperature: its thermal optimum varies from 29°C in early spring to 41°C in summer and so to 28°C in autumn

Figure 6. Encelia sp. (Yellow flowers) is a typical constitute of dry out areas in California (here Palm Canyon trail). [Source: © Stan Shebs, via Wikimedia commons CC Past-SA 3.0]

Changes in the thermal optimum can be even more rapid and of great amplitude. For example, in a seaside clone [5] of Encelia californica, a change in growth temperature from 30°C (constant day and night temperature) to fifteen°C during the day and 2°C during the night for three days is sufficient to lower the thermal optimum past about x degrees.

In general, these changes tin can be measured in both growing and mature leaves, with the response being of greater amplitude in growing leaves.

2.5. Heat-sensitive versuscold-sensitive species

Warm acclimation of cool-adapted species (or ecotypes [vi]) occurs with an increase in thermal optimum but a general subtract in photosynthesis.

This is, for case, the case ofAtriplex sabulosa. One tin can then wonder about the interest of this change. The reverse may be truthful for plants strictly adapted to warm conditions, such equally Tridestomia oblongifolia. Figure 7 illustrates the case ofAtripex lentiformis, [vii] a perennial leafy plant, which occurs in California in both Decease Valley and in cool, wet coastal habitats:

• The assimilation of the desert ecotype (Figure 7A) and the coastal ecotype (Figure 7B) show almost the same response to temperature when grown under 23°C during the day and 18°C at nighttime (in ruby-red in Figure 7).

• 

  Figure 7. Variations in CO2 assimilation of Atriplex lentiformis ecotypes from a warm surround (A) and a cool, moist environment (B). [Source: Author'due south diagram, after Pearcy (1977)] Correct, Atripex lentiformis (salt bush) [Source: Forest & Kim Starr, CC BY 3.0, via Wikimedia Eatables]

  Under the alternating 43°C day and xxx°C nighttime (blue in Effigy 7), only the desert ecotype shows plasticity, maintaining high CO_two assimilation under these new conditions. The activity of the coastal ecotype is low at all temperatures. Only the displacement of the thermal optimum remains of its acclimatization capabilities [8].

ii.6. C3 plants versus C4 plants

Figure eight. Chief forest in southern Argentina. Almost all copse are C3 plants. [Source: © G. Cornic]

C3 plants were the first to appear and constitute about 85% of current plant species. They mainly colonize absurd and humid environments (or seasons). Trees, for example, with rare exceptions, are C3 plants (Read The path of carbon in photosynthesis) (Figure 8).

C4 plants, of which there are traces only from the finish of the Tertiary Era, constitute only 5% of the species. They tend to colonize hot and dry environments (or seasons) (See Restoring savannas and tropical herbaceous ecosystems). Maize and sugarcane are examples.

On average, the thermal optimum of C4 plants is located at higher temperatures than that of C3 plants.

However, C3 plants are the nigh plastic. In fact, their thermal optimum varies from around seven to 35 ° C, while that of C4 plants oscillates, with a few exceptions, between thirty and xl ° C. In add-on, when the temperature is beneath xx ° C, the photosynthesis of C4 plants is on average lower than that of C3 plants.

3. CO₂ absorption results from the interaction of processes whose response to temperature is different

The absorption of calorie-free at the collecting antennae (Figure ix) and the transfer of its free energy to the PSII reaction centres are non temperature sensitive.
Are temperature sensitive:

• The diffusion of CO_two  from the ambience air to the chloroplasts: its speed increases with temperature.
• The fixation of CO₂  on Ribulose 1,five-bisphosphate (RuBP), a sugar whose skeleton is formed past 5 carbon atoms (Read Focus Deciphering the Benson-Bassham-Calvin cycle)
• The transfer of electrons from PSII to PSI.

Figure 9. Diagram of the interacting processes during photosynthetic CO2 fixation (case of a C3 plant). PSI and PSII: respectively photosystem I and Ii. They are included in the thylakoid membrane, which is fabricated up of two lipid layers forming "sacs" in the chloroplast. The interior of the thylakoid is the lumen. RubisCO: enzyme that catalyzes the fixation of CO2 on a carbohydrate with 5 carbon atoms (Ribulose 1,five-bisphosphate: C5). Benson-Calvin bike: allows the regeneration of C5, and at the same time gives the institute the necessary carbon. ATP is synthesized when protons from the lumen return to the stroma through an ATPase using inorganic phosphate, Pi. The lumen protons have ii origins: (i) oxidation of water in the lumen past PSII which also provides electrons, e- and (2) operation of a proton pump in the thylakoid that passes protons from the stroma into the lumen. [Source: Author's diagram]

The regeneration of RuBP occurs via the operation of the Benson-Calvin cycle (This is the "biochemistry" of the procedure) which uses reducing ability (in the course of NADPH) provided by electron transfer to function. The necessary ATP is synthesized when protons accumulated in the lumen pass into the stroma through an ATPase (Figure 9).
The formation of reducing power and the synthesis of ATP have a thermal sensitivity close to that of electron transfer.

4. What are the processes at work in setting the thermal optimum for CO_two assimilation in C3 and C4 plants?

four.ane. Photosystem action and the resulting electron transfer are non involved

Measured in vitro on isolated thylakoids (see legend Figure ix), in the presence of artificial acceptors, electron transfer increases with temperature and shows a clear thermal optimum. It is located around xxx°C and corresponds to that of CO_two assimilation when the latter is saturating [ix]. The activity of PSII has a thermal optimum identical to that of the electron transfer chain.

• PSI action is not inhibited at high temperatures (higher up 30°C, upwardly to 45°C) where it remains stable or even increases: information technology is the activity of PSII that limits the activity of the electron chain.
• Moreover, PSII is very sensitive to high temperatures which damage the protein circuitous that allows the oxidation of water (encounter Figure 9).

The thermal response of electron transfer is like in C3 and C4 plants. However, there are organizational differences betwixt these two types of plants (see The path of carbon in photosynthesis).
The supply of energy cannot therefore explicate the differences in thermal optimum. It is the way in which the energy produced is used that makes the difference.

4.2. An answer? Comparison of the effect of atmospheric O₂ on CO₂ assimilation of C3 and C4 plants

• In normal air [10], 21% O₂ (+ North₂) + 360 ppm CO₂: the thermal optimum is 27°C in Maize (C4 found), while it is only 22°C in Pea (C3 plant) (Figure ten): thethermal optimum of the C4 plant is higher than that of the C3 plant (see besides section 2.6).
• In an oxygen-scarce atmosphere, 1% O₂ (+ N₂) + 360 ppm CO_two: the CO₂ uptake of Maize is not affected, while that of Pea is stimulated above almost 17°C, with a shift in its thermal optimum to near that of Maize.
• In C3 plants, atmospheric oxygen inhibits CO_ii  uptake when the leaf temperature is sufficiently high, whereas information technology has no result (or negligible effect) in C3 plants

• 

  Figure 10. Variation of CO2 absorption measured in leaves of Pea (A; Pisum sativum) and Maize (B; Zea mays) equally a function of leaf temperature. The plants were grown in natural calorie-free at a temperature of twenty ± 2°C. [Source: Author's diagram – royalty-free epitome / Pixabay]

  Note that the variation in electron transfer estimated in vivo, by measuring chlorophyll fluorescence emission as a role of temperature, is very similar in 1% and 21% O₂ in Pea: the variation in thermal optimum is therefore non due to a alter in photochemistry.

4.3. Rubisco properties explain the departure in response

• Case of C3 plants

CO₂and O₂compete to occupy the active sites of Rubisco: This enzyme has a carboxylase function and an oxygenase function. CO₂ enters the Benson-Calvin bike and the photosynthetic fixation of O₂ is at the origin of a metabolic pathway responsible for photorespiration (Effigy 11; see also The path of carbon in photosynthesis).

CO_two occupies a high number of active sites on the Rubisco when the O₂ content of the ambience air is low (1% for example) or that of CO₂is high.

O₂ is mainly fixed if its content increases or if that of CO₂  decreases (the latter then releases active sites which are then occupied past O_{ii )}.

In normal air, there are ii reasons why O₂ fixation increases (and consequently CO₂ fixation decreases) when the temperature increases [xi].

1. The affinity of Rubisco for_CO2 decreases more than that for O₂; a factor that favours the assimilation of O_ii.
2. The water solubility coefficient of CO₂  decreases more than than that of O_ii, leading to a more rapid decrease in the amount of CO₂ than O₂ in the chloroplast; this is a gene that favours O₂ fixation.

In an O_two-poor atmosphere(Figure ten), competition between O₂ and CO₂is very reduced. Energy is so used mainly for CO₂ assimilation, which increases in value until around thirty°C and and so decreases as the energy supply decreases (meet section 4.1).

In normal air, the consequence of O_two on photosynthetic CO₂ fixation (Figure xi) is very depression (or fifty-fifty nil) when the temperature is low: competition on the carboxylation sites is in favour of CO_2.

Figure 11. Schematic of CO2 and O2 fixation on RuBP (Ribulose 1,v-bisphosphate) in a C3 plant. APG: 3-phosphoglyceric acid, 3 C chemical compound; TP: Trioses phosphate. The carbon leaves the Calvin bicycle to feed the synthesis of sucrose. PG: phosphoglycolate, 2C compound. Two PGs give a serine (Ser) containing 3C with the product of CO2 from photorespiration. C = Carbon atom. Source: Author's diagram]

On the other hand, when the temperature increases, the competition on these sites favours the fixation of O_ii which then consumes an increasing role of the energy produced by the activity of the photosystems. This energy is therefore no longer available for CO₂ fixation, which reaches its maximum value around 22°C.

• Case of C4 plants.

CO_iiis concentrated at the Rubisco by a mechanism that is insensitive to oxygen. Its content can reach 800 to 2000 ppm depending on the plant in C4: that is to say contents from 2 to 5 times higher than its electric current atmospheric content.

Under these conditions, photosynthetic O₂ fixation is weak or even non-existent because the active sites of the Rubisco are all occupied by CO₂. The energy supplied by the activity of the photosystems is therefore used only in the fixation of CO₂  when the leaf temperature increases, explaining the college thermal optimum in this blazon of constitute.

C4 plants evolved from C3 plants during the global subtract in atmospheric CO_two content at the end of the 3rd Era [12].

This decrease would then have "released" the oxygenase function of the Rubisco of C3 plants, resulting in a loss of fixed carbon via photorespiration.

The institution of a CO₂ concentration mechanism is an advantage because it prevents this carbon loss. Nosotros currently find species that are "intermediates" between C3 and C4.

5. The thermal optimum of C3 photosynthesis is modulated by certain environmental parameters

5.ane. The CO₂ content in the atmosphere

The thermal optimum increases with increasing ambient CO₂ content. In the case shown in Figure 12, it increases from virtually 10°C when the content is 100 ppm to more than 30°C when it is 800 ppm.

Effigy 12. Variations in CO2 uptake equally a role of leaf temperature measured on a Pea leaf placed at different ambient CO2 levels. Light near saturation. [Source: © Chiliad. Cornic, unpublished]

This effect is explained by the competition between CO₂  and O_ii for the occupation of the active sites of the Rubisco: at 800 ppm CO_ii the active sites are occupied mainly by CO₂ ; at 100 ppm CO_ii the occupation of these sites past atmospheric O₂ is in majority.

Effigy 13. Human activities lead to an increment in carbon dioxide in the atmosphere. Its content went from 320 to 415 ppm in the space of 50 years. This increase has consequences on the temperature of the atmosphere and the action of the vegetation. [Source : Royalty-costless image / Pixabay]

 In a globe with steadily increasing atmospheric CO₂ (Figure xiii), the thermal optimum of C3 plants is expected to increase. This does not hateful, even so, that plant product will then be higher (encounter note 3 section one): episodes of high heat will, like droughts, certainly be more frequent.

5.ii. Lack of water

The photosynthetic apparatus is resistant to drought. Information technology retains all its capacity to absorb CO_two on the Rubisco, and to produce free energy until the leaves have lost about xxx% of their water [13].

• CO_ii uptake decreases in this range of water loss, because the stomata shut (meet Focus Leaf transpiration and heat protection). This closure slows down the entry of CO₂into the leaf and consequently leads to a decrease of the CO_ii  content in the mesophyll.
• Yet, the O_two content in the chloroplasts remains high. Indeed, its content in the atmosphere (21% or 210,000 ppm) is, compared to that of CO_ii (@ 400 ppm), very loftier and in any case sufficient for a very substantial quantity to pass through the epidermis fifty-fifty when the stomata are closed.
• The competition betwixt CO₂ and O₂ for the occupation of the active sites of the Rubisco is thus in favour of O₂ .

Effigy 14. A, Variations in CO2 assimilation as a part of leaf temperature. Leaves with different amounts of water loss found in air with an ambient CO2 content of 400 ppm. B, The electron transfer charge per unit estimated on the aforementioned leaves by measuring the chlorophyll fluorescence emission. [Source: Author's diagram, afterwards Cornic et al. ref. 14]

Therefore, the thermal optimum for photosynthesis must lower in C3 plants that dry out.

This is shown in Effigy 14A, in which the thermal optimum drops from about 23°C, in a Pea leafage at maximum turgor, to 17°C when it has lost 20% of its h2o.

Electron transfer in the thylakoid membrane is not affected by water loss in the range shown (Figure 14B). When water loss is 20%, the free energy produced by photosystem activity is primarily used to bind atmospheric oxygen to RuBP [14], resulting in increased photorespiration.

vi. Why, from its thermal optimum, CO₂ absorption decreases as temperature decreases or increases?

6.1. When the temperature lowers

Several reasons probably all contribute, to varying degrees, to this decrease :

• The rate of RuBP turnover decreases: there is a slowdown in the activeness of some enzymes decision-making this turnover, notably that of a Fructose 1,half-dozen-bisphosphate (run into Figures 9 and eleven).
• Sequestration of phosphorylated compounds in chloroplasts. The triose phosphate is no longer (or less) exported when sucrose synthesis is inhibited. The inorganic phosphate in the chloroplast is no longer renewed leading to a subtract in ATP synthesis.
• Inhibition of the electron transfer concatenation (run into department 4.one), resulting in reduced energy production (reducing power and ATP).

In C4 plants information technology is the activity of the Rubisco that appears to be preponderant, although the cold sensitivity of enzymes involved in CO₂ aggregating at the Rubisco is well known.

6.2. As the temperature increases

In C3 plants the increase in photorespiration decreases the fraction of electrons produced by PSII and used to assimilate CO₂. Nevertheless, other factors are at play since CO_two assimilation measured (1) in an atmosphere with little or no photorespiration (ambient O₂ content of i%), and (ii) measured in a normal atmosphere in a C4 establish decreases in both cases (Figure 10).

Several reasons can exist given:

• The slowing down of PSII action leading to that of the electron transfer chain from PSII to PSI.
• To perform its part Rubisco must exist activated past an enzyme called Rubisco activase, the activity of which decreases when the temperature is college than near 33°C (incidentally, high-temperature resistant activases appear in some plants subjected to periods of loftier heat [15]). Notwithstanding, since activase must itself be activated past an electron transfer-dependent process, information technology cannot exist ruled out that the latter is besides involved in limiting [15].
• The "catalytic misfiring" of Rubisco increases with temperature and increasing amounts of an inhibitor of the enzyme (Xylulose-1,4-bisphosphate), which is structurally close to RuBP (encounter Figures 9 and 11), are synthesized.

In C4 plants(case of Maize) the activation and activeness of enzymes that participate in the CO₂ concentration organisation at the Rubisco are not very sensitive to high temperatures. The same reasons as in a higher place may explain the decrease in CO₂assimilation when the temperature increases across that of the thermal optimum.

7. Hardening after constitute exposure to absurd (≤ about ten°C) and high (≥ about 37°C) temperatures

Maintaining plants at cool or loftier temperatures causes, forth with the changes in photosynthesis described in a higher place, increment in their resistance to otherwise lethal temperatures(frost and high temperature). This is hardening.

In this process, temperature and lite interact and the metabolic changes induced are sometimes very rapid (from minutes to hours).

Thus, cold hardening can exist accomplished at ordinary temperature past modulating the length of the low-cal period or its spectral composition in the red [16]. All the same, cold is still required to attain full hardening. Too the lack of low-cal in the cold prevents hardening to varying degrees.

• At elevated temperatures : the transmitted signals activate the synthesis of chaperone proteins (HSPs: Heat Schock Proteins) that repair denaturing proteins, also forestall their coagulation or even help marking them for degradation.
• At absurd temperatures: the synthesis of chaperone proteins is also activated. It is accompanied by (i) the synthesis of "antifreeze" proteins that interfere with ice crystal germination and (ii) an increase in sugar synthesis disposed to increment osmotic pressure in the cells.

Note that the signaling pathways and their interactions inducing the genome response are but partially known. The references given in "Larn More" and an fastened Focus permit for further exploration of this evolving point.

8. Effects of temperature on photosynthesis: summary diagram

The summary diagram (Figure fifteen) classifies the furnishings of temperature on photosynthesis according to the speed of temperature change and the extent of its variation. Note that hardening allows leafage maintenance in perennial leafage plants and therefore minimizes energy loss under extreme temperature weather condition.

Figure xv. Scheme classifying the furnishings of temperature on photosynthesis. [Source: Author'southward diagram]

The rapidity of electric current climatic change makes it necessary to delve deeper into the responses of plants to their environment: the hope is to be able to maintain sufficient primary production to keep the biosphere performance.

ix. Messages to remember

• The uptake of CO₂ by a leaf has a thermal optimum close to the average temperature of its growth environment.
• This thermal optimum tin change rapidly when the conditions of the environment are durably modified: this is a process acclimatization.
• This thermal optimum is on average less in C3 plants than in C4 plants: this is mainly due to photosynthetic fixation of atmospheric O₂ via Rubisco action in C3 plants.
• This optimum depends on the CO₂ content of the ambient air in C3 plants: at high content it becomes identical to that in C4 plants.
• This optimum depends on the hydration state of the leaf.
• Subjected to absurd or hot temperatures plants bring into play processes hardening to otherwise lethal temperatures. These processes involve protein syntheses and changes in the fluidity of chloroplast and cell membranes.

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Notes and references

Cover image. Sunset over the Sonora Arizona desert. [Source: royalty free / Pixabay]

[1] Meehl GA, Stocker TF, Collins WD, Riedlingstein P, Gaye AT, Gregory JM, Kitoh A, Knutti R, Murphy JM, Noda A & Raper SCB (2007). Climatic change 2007: The Physical Scientific discipline Ground. Contribution of Working Group I to the Quaternary Assessment Report of the Intergovernmental Console on Climate Alter. Cambridge University Press

[two] Yamori W, Hikosaka K & Manner DA. (2014). Temperature response of photosynthesis in C3, C4, and CAM plants: temperature acclimation and temperature accommodation. Photosynthesis Res. 119, 101- 117.

[iii] For example, when growing plants are subjected to drought, the amount of carbon they assimilate decreases initially because leaf growth is inhibited. The mechanisms for CO_two fixation in the foliage are non so inhibited. Boyer JS (1970) Institute Physiol.46, 233-235

[4] The values of thermal optima given here, are from measurements fabricated in "normal air", containing 21% O_two and about 400 ppm CO₂. When this is non the case the O₂ and CO₂ contents are shown. The CO₂ uptake in air containing 21% O₂ is saturated from nigh 1200 ppm CO₂ when light is close to saturation. The evaporative ability of the air is also regulated in almost cases during the measurements. It is estimated by the saturation deficit of the fractional pressure of h2o vapor in the ambient air effectually the leaves.

[5] Plants from the same individual by vegetative reproduction. They are genetically identical.

[6] Ecotype: Plants of the same species from dissimilar environments, which, grown from seed to blossom under identical weather condition prove unlike physiological characteristics.

[vii] It fetches h2o from as far equally the water table, hence its name of phreatophyte plant.

[8] Pearcy RW (1971). Acclimation of photosynthetic and respiratory CO₂ exchange to growth temperature in Atriplex lentiJormis (Torr.) Wats. Establish Physiol. 59, 795-799

[9] Yamasaki T, Yamakawa T, Yamane Y, koike H, Satoh M & Katoh S. (2002) Temperature acclimation of photosynthesis and related changes in photosystem II electron transport in winter wheat. Plant Physiol. 128 1087-1097.

[10] See annotation #4, section two.2

[11] Jordan DB & Ogren WL (1984). The CO₂/O₂ specificity of ribulose 1,5-bisphosphate carboxylase/oxygenase. Dependence on ribulose bisphosphate concentration, pH and temperature. Planta 161, 308-313

[12] Ehleringer JR, Sage RF, Flanagan LB & Pearcy RW (1991). Climate change and the evolution of C4 photosynthesis. Trends in Ecology and Evolution half-dozen, 95-99

[13] Cornic G & Massacci A (1996). Leaf photosynthesis under drought stress. In Advances in Photosynthesis (vol v) Photosynthesis and the surround, 347-366. Neil R Bakery (ed.) Kluwer Bookish publishers Dordrecht.

[14] Cornic One thousand, Badeck F-W, Ghashghaie J & Manuel N (1999). Issue of temperature on net CO₂ uptake, stomatal conductance for CO₂ and quantum yield of photosystem Ii photochemistry of dehydrated pea leaves. In Sanchez Dias M, Irigoyen JJ, Aguirreolea J & Pithan K (eds) Ingather development for absurd and moisture regions of Europe. European community. ISBN 92-828-6947-4.

[15] Crafts-Brandner SJ, van de Loo FJ & Salvucci ME (1997). The two forms of ribulose-1,v-bisphosphate carboxylase/oxygenase activase differ in sensitivity to elevated temperature. Plant Physiol. 114, 439-444.

[xvi] Puhakainen T, Li C, Boije-Malm M, Kangasjärvi J, Heino P & Palva ET. (2004). Brusque-24-hour interval potentiation of depression temperature-induced gene expression of a C-echo-binding factor-controlled gene during cold acclimation in Argent Birch. Establish Physiol.136, 4299-4307

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The Encyclopedia of the Surroundings by the Association des Encyclopédies de fifty'Environnement et de l'Énergie (www.a3e.fr), contractually linked to the University of Grenoble Alpes and Grenoble INP, and sponsored by the French Academy of Sciences.

To cite this commodity: CORNIC Gabriel (2022), Effects of temperature on photosynthesis, Encyclopedia of the Environment, [online ISSN 2555-0950] url : https://world wide web.encyclopedie-environnement.org/en/life/effects-temperature-on-photosynthesis/.

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