Terrestrial plants are regularly subjected to strong temperature variations. These variations tin reach an amplitude of twoscore°C or even more, both in polar regions and in hot desert areas. Beingness rooted, they take reduced mobility and must cope with changes in their surround. The assimilation of CO2 by plants via photosynthesis is the gateway to carbon in the biosphere. What is the thermal amplitude that allows it to function? How does photosynthesis reacts to rapid and dull temperature variations? What is the diversity of responses? What are the physiological processes that limit it? Crucial questions tare o be considered in the context of global warming.
one. Institute product and climate change
The electric current increase in greenhouse gas emissions will cause an increase in atmospheric temperature of 2 to 3°C in the adjacent 50 years (run across A carbon wheel disrupted past human activities). At the same time, heat waves and extreme heat periods volition be more than frequent and of longer duration [one]. Agricultural production and the functioning of forests volition therefore be greatly affected. Models based on large-scale observations point that, in the absenteeism of agronomic adaptation, the subtract in crop yields tin can reach 17% for each i°C increase in the temperature of the growing flavour [two].
The production of college plants depends in particular (just not just [3]) on leaf photosynthesis (come across Shedding light on photosynthesis & The The path of carbon in photosynthesis). COii enters the leafage where its reduction in the chloroplasts is accompanied by O2 production. Its entry is almost exclusively through the stomata (Effigy 1). For each molecule of CO2 absorbed, l to 300 molecules of water are transpired from the leaves, depending on the plant. This water allows, among other things, the cooling of the leaf (see Focus Foliage transpiration and heat protection).
Figure ane. During photosynthesis, CO2 is captivated and O2 is released mainly through the stomatal opening (ostiole). The h2o vapour (transpiration of the leaf) passes mainly through the ostiole but too through the epidermis. Transpiration allows the leaf to absurd down in the light. [Source: Writer'due south diagram]
The leaf is a converter of solar free energy into chemical energy and, like any energy converter, requires a permanent cooling organization.
The climate changes that are currently occurring make it necessary to empathize the effects of temperature on photosynthesis.
two. The thermal optimum of photosynthesis
two.1. Diagram of the thermal response
Photosynthetic CO2 uptake varies with temperature. In most cases its response to temperature is rapidly reversible between nearly 10 and 34°C. In this range of temperatures information technology presents a maximum value: a thermal optimum.
Figure two. 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 10°C and higher up 34°C plants get-go to gear up protective mechanisms. For these extreme values, COtwo absorption is oftentimes unstable and tin be cancelled more or less speedily: the foliage is then under stress (Figure 2).
2.ii. A thermal optimum based on the average temperature of the surroundings
Plants in common cold environments or with a cold growing season have a higher photosynthesis at low temperatures. Plants in warm environments, or growing during the warm season, have a higher photosynthesis at loftier temperatures.
Figure 3. Deschampsia antarctica is one of two flowering plants found in Antarctica. Information technology is oft subjected to negative temperatures. The snow that frequently covers it protects it from extreme temperatures. [Source: Lomvi2, CC By-SA 3.0, via Wikimedia Eatables]
For instance, the thermal optimum for COtwoassimilation [4] in Deschampsia antarctica (Figure 3) and Colobanthus quitensis, the only two Antarctic flowering plants, is between 8 and fifteen°C, while it is effectually 45°C in Tridestomia oblongifolia, a warm desert plant from Central America. The latter species probably holds the world record for flowering plants in this respect.
2.three. Acclimatization to the thermal conditions of the environment
Figure 4. Variations in CO2 absorption as a function of leaf temperature, in a constitute grown at x°C (red) or 25°C. Measurements fabricated on Pea, under a calorie-free close to saturation. CO2 content in ambient air: 390 ppm. [Source: Writer's diagram]
Differences in the thermal response of photosynthesis are besides found in individuals of the same species growing at unlike temperatures. Figure 4 shows COtwo assimilation in pea grown at ten or 25°C.
In the outset case (cultivation at x°C) the thermal optimum is nigh 16°C, while it is higher than 25°C in the second (tillage at 25°C). At low temperatures, CO2 absorption is college in plants grown at x°C.
In this instance the adjustment to absurd atmospheric condition is a gain for the institute.
2.4. Acclimatization can exist rapid
Effigy v. Remth (Hammada scoparia), a characteristic plant of the Wadi Rum desert (Jordan). [Source: Ji-Elle, CC Past-SA 4.0, via Wikimedia Commons]
For example, the photosynthesis of Hammada scoparia, a bush in the deserts of the Middle East (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 then to 28°C in autumn
Changes in the thermal optimum can be fifty-fifty more rapid and of great amplitude. For example, in a seaside clone [5] of Encelia californica, a change in growth temperature from 30°C (abiding day and night temperature) to fifteen°C during the day and 2°C during the night for 3 days is sufficient to lower the thermal optimum by near 10 degrees.
In full general, these changes can exist 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 [6]) occurs with an increase in thermal optimum simply a general decrease in photosynthesis.
This is, for example, the example ofAtriplex sabulosa. One can then wonder nigh the involvement of this alter. The opposite may be true for plants strictly adjusted to warm conditions, such as Tridestomia oblongifolia. Figure vii illustrates the case ofAtripex lentiformis, [7] a perennial leafy found, which occurs in California in both Death Valley and in cool, wet littoral habitats:
The assimilation of the desert ecotype (Effigy 7A) and the coastal ecotype (Figure 7B) show almost the same response to temperature when grown under 23°C during the day and xviii°C at night (in red in Effigy 7).
Effigy vii. Variations in CO2 absorption of Atriplex lentiformis ecotypes from a warm environment (A) and a cool, moist environs (B). [Source: Author's diagram, after Pearcy (1977)] Correct, Atripex lentiformis (salt bush) [Source: Forest & Kim Starr, CC BY 3.0, via Wikimedia Eatables]Under the alternate 43°C day and 30°C nighttime (blueish in Effigy vii), simply the desert ecotype shows plasticity, maintaining high CO2 assimilation under these new atmospheric condition. The action of the coastal ecotype is low at all temperatures. Only the displacement of the thermal optimum remains of its acclimatization capabilities [8].
C3 plants were the outset to announced and institute 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 in that location are traces merely from the terminate of the 3rd 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 boilerplate, the thermal optimum of C4 plants is located at higher temperatures than that of C3 plants.
Withal, C3 plants are the most plastic. In fact, their thermal optimum varies from around 7 to 35 ° C, while that of C4 plants oscillates, with a few exceptions, between 30 and 40 ° C. In addition, when the temperature is below twenty ° C, the photosynthesis of C4 plants is on average lower than that of C3 plants.
three. CO2 absorption results from the interaction of processes whose response to temperature is unlike
The absorption of light at the collecting antennae (Figure 9) and the transfer of its energy to the PSII reaction centres are not temperature sensitive. Are temperature sensitive:
The improvidence of CO2 from the ambience air to the chloroplasts: its speed increases with temperature.
The fixation of CO2 on Ribulose ane,5-bisphosphate (RuBP), a carbohydrate whose skeleton is formed by 5 carbon atoms (Read Focus Deciphering the Benson-Bassham-Calvin cycle)
The transfer of electrons from PSII to PSI.
Effigy 9. Diagram of the interacting processes during photosynthetic CO2 fixation (case of a C3 plant). PSI and PSII: respectively photosystem I and 2. They are included in the thylakoid membrane, which is made up of ii 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 sugar with 5 carbon atoms (Ribulose 1,5-bisphosphate: C5). Benson-Calvin wheel: allows the regeneration of C5, and at the same fourth dimension gives the plant 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: (1) oxidation of h2o in the lumen by PSII which also provides electrons, e- and (ii) 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 functioning of the Benson-Calvin bicycle (This is the "biochemistry" of the process) which uses reducing power (in the grade of NADPH) provided by electron transfer to role. The necessary ATP is synthesized when protons accumulated in the lumen pass into the stroma through an ATPase (Effigy nine). The germination of reducing power and the synthesis of ATP take a thermal sensitivity close to that of electron transfer.
4. What are the processes at work in setting the thermal optimum for CO2 assimilation in C3 and C4 plants?
4.1. Photosystem activity and the resulting electron transfer are not involved
Measured in vitro on isolated thylakoids (see legend Figure 9), in the presence of artificial acceptors, electron transfer increases with temperature and shows a clear thermal optimum. It is located around 30°C and corresponds to that of CO2 absorption when the latter is saturating [ix]. The action of PSII has a thermal optimum identical to that of the electron transfer chain.
PSI action is not inhibited at high temperatures (above thirty°C, up to 45°C) where it remains stable or fifty-fifty increases: it is the action of PSII that limits the activeness of the electron chain.
Moreover, PSII is very sensitive to loftier temperatures which impairment the poly peptide complex that allows the oxidation of water (see Figure ix).
The thermal response of electron transfer is similar in C3 and C4 plants. Notwithstanding, there are organizational differences between these 2 types of plants (meet The path of carbon in photosynthesis). The supply of energy cannot therefore explicate the differences in thermal optimum. It is the style in which the energy produced is used that makes the deviation.
4.two. An answer? Comparison of the effect of atmospheric O2 on CO2 assimilation of C3 and C4 plants
In normal air [x], 21% O2 (+ N2) + 360 ppm COii: the thermal optimum is 27°C in Maize (C4 plant), while it is only 22°C in Pea (C3 institute) (Figure 10): thethermal optimum of the C4 plant is higher than that of the C3 plant (see also department two.vi).
In an oxygen-scarce atmosphere, 1% O2 (+ N2) + 360 ppm COii: the COtwouptake of Maize is not affected, while that of Pea is stimulated above near 17°C, with a shift in its thermal optimum to well-nigh that of Maize.
In C3 plants, atmospheric oxygen inhibits COii uptake when the leaf temperature is sufficiently high, whereas it has no upshot (or negligible result) in C3 plants
Figure 10. Variation of CO2 assimilation measured in leaves of Pea (A; Pisum sativum) and Maize (B; Zea mays) every bit a function of leaf temperature. The plants were grown in natural light at a temperature of twenty ± two°C. [Source: Writer's diagram – royalty-free image / Pixabay]Note that the variation in electron transfer estimated in vivo, past measuring chlorophyll fluorescence emission every bit a role of temperature, is very like in 1% and 21% O2 in Pea: the variation in thermal optimum is therefore not due to a modify in photochemistry.
4.3. Rubisco properties explain the difference in response
Instance of C3 plants
CO2and O2compete to occupy the active sites of Rubisco: This enzyme has a carboxylase function and an oxygenase function. COtwo enters the Benson-Calvin wheel and the photosynthetic fixation of Oiiis at the origin of a metabolic pathway responsible for photorespiration (Figure 11; come across likewise The path of carbon in photosynthesis).
COii occupies a high number of active sites on the Rubisco when the O2 content of the ambience air is low (one% for case) or that of CO2is loftier.
O2 is mainly stock-still if its content increasesor if that of COtwo decreases (the latter then releases active sites which are then occupied past Oii).
In normal air, in that location are 2 reasons why O2 fixation increases (and consequently CO2 fixation decreases) when the temperature increases [11].
The affinity of Rubisco forCO2decreases more than that for Otwo; a factor that favours the absorption of O2.
The water solubility coefficient of CO2 decreases more than that of O2, leading to a more than rapid decrease in the amount of CO2 than O2 in the chloroplast; this is a gene that favours Otwo fixation.
In an O2-poor temper(Figure x), contest between O2 and COtwois very reduced. Energy is then used mainly for COtwo absorption, which increases in value until around 30°C and then decreases as the energy supply decreases (see section 4.ane).
In normal air, the outcome of O2 on photosynthetic COii fixation (Effigy eleven) is very low (or even nothing) when the temperature is low: contest on the carboxylation sites is in favour of CO2.
Effigy 11. Schematic of CO2 and O2 fixation on RuBP (Ribulose 1,v-bisphosphate) in a C3 plant. APG: three-phosphoglyceric acid, 3 C compound; TP: Trioses phosphate. The carbon leaves the Calvin bike to feed the synthesis of sucrose. PG: phosphoglycolate, 2C compound. Ii PGs give a serine (Ser) containing 3C with the production 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 O2which so consumes an increasing role of the energy produced past the activity of the photosystems. This energy is therefore no longer available for COtwo fixation, which reaches its maximum value around 22°C.
Case of C4 plants.
COtwois concentrated at the Rubisco by a mechanism that is insensitive to oxygen. Its content can accomplish 800 to 2000 ppm depending on the plant in C4: that is to say contents from 2 to 5 times college than its electric current atmospheric content.
Under these conditions, photosynthetic O2 fixation is weak or even non-real considering the active sites of the Rubisco are all occupied past CO2. The energy supplied by the activity of the photosystems is therefore used simply in the fixation of COii when the leaf temperature increases, explaining the college thermal optimum in this type of plant.
C4 plants evolved from C3 plants during the global decrease in atmospheric CO2content at the terminate of the Tertiary Era [12].
This decrease would and then have "released" the oxygenase function of the Rubisco of C3 plants, resulting in a loss of fixed carbon via photorespiration.
The establishment of a COii concentration mechanism is an advantage considering it prevents this carbon loss. We currently find species that are "intermediates" between C3 and C4.
5. The thermal optimum of C3 photosynthesis is modulated by certain ecology parameters
5.1. The COiicontent in the temper
The thermal optimum increases with increasing ambient CO2 content. In the case shown in Effigy 12, it increases from about 10°C when the content is 100 ppm to more than xxx°C when it is 800 ppm.
This effect is explained by the competition between CO2 and O2 for the occupation of the active sites of the Rubisco: at 800 ppm CO2 the agile sites are occupied mainly past CO2 ; at 100 ppm COtwo the occupation of these sites past atmospheric O2 is in bulk.
Figure thirteen. Human activities atomic number 82 to an increase in carbon dioxide in the temper. 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-free image / Pixabay]
In a world with steadily increasingatmospheric CO2(Figure 13), the thermal optimum of C3 plants is expected to increment. This does non hateful, yet, that plant production volition and then be higher (run into note iii section i): episodes of high heat will, like droughts, certainly be more frequent.
five.2. Lack of water
The photosynthetic apparatus is resistant to drought. Information technology retains all its chapters to absorb CO2 on the Rubisco, and to produce energy until the leaves have lost nearly 30% of their water [13].
CO2 uptake decreases in this range of water loss, because the stomata shut (encounter Focus Leaf transpiration and oestrus protection). This closure slows downwardly the entry of CO2into the leaf and consequently leads to a decrease of the CO2 content in the mesophyll.
Even so, the Otwo content in the chloroplasts remains high. Indeed, its content in the atmosphere (21% or 210,000 ppm) is, compared to that of COtwo (@ 400 ppm), very high and in whatever case sufficient for a very substantial quantity to pass through the epidermis even when the stomata are closed.
The contest betwixt COtwo and O2 for the occupation of the agile sites of the Rubisco is thus in favour of O2.
Figure 14. A, Variations in CO2 assimilation as a part of leafage temperature. Leaves with unlike amounts of water loss constitute in air with an ambient CO2 content of 400 ppm. B, The electron transfer rate estimated on the aforementioned leaves by measuring the chlorophyll fluorescence emission. [Source: Author'southward diagram, after Cornic et al. ref. xiv]
Therefore, the thermal optimum for photosynthesis must lower in C3 plants that dry out out.
This is shown in Effigy 14A, in which the thermal optimum drops from nearly 23°C, in a Pea leaf at maximum turgor, to 17°C when it has lost 20% of its water.
Electron transfer in the thylakoid membrane is not affected past h2o loss in the range shown (Figure 14B). When water loss is twenty%, the energy produced by photosystem activity is primarily used to bind atmospheric oxygen to RuBP [14], resulting in increased photorespiration.
6. Why, from its thermal optimum, COtwo assimilation decreases as temperature decreases or increases?
6.i. When the temperature lowers
Several reasons probably all contribute, to varying degrees, to this subtract :
The rate of RuBP turnover decreases: there is a slowdown in the activity of some enzymes decision-making this turnover, notably that of a Fructose 1,6-bisphosphate (encounter Figures ix 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 decrease in ATP synthesis.
Inhibition of the electron transfer chain (come across department 4.one), resulting in reduced energy production (reducing ability and ATP).
In C4 plants it is the activity of the Rubisco that appears to be preponderant, although the cold sensitivity of enzymes involved in CO2 accumulation at the Rubisco is well known.
6.2. As the temperature increases
In C3 plants the increment in photorespiration decreases the fraction of electrons produced by PSII and used to assimilate CO2. Even so, other factors are at play since COii assimilation measured (1) in an atmosphere with little or no photorespiration (ambience Oii content of i%), and (two) measured in a normal atmosphere in a C4 plant decreases in both cases (Effigy ten).
Several reasons can be given:
The slowing downwardly of PSII action leading to that of the electron transfer concatenation from PSII to PSI.
To perform its role Rubisco must exist activated by an enzyme called Rubisco activase, the activity of which decreases when the temperature is higher than about 33°C (incidentally, high-temperature resistant activases announced in some plants subjected to periods of loftier heat [15]). However, since activase must itself be activated by an electron transfer-dependent process, it cannot be ruled out that the latter is also involved in limiting [15].
The "catalytic misfiring" of Rubisco increases with temperature and increasing amounts of an inhibitor of the enzyme (Xylulose-1,four-bisphosphate), which is structurally close to RuBP (run into Figures 9 and 11), are synthesized.
In C4 plants(case of Maize) the activation and activity of enzymes that participate in the CO2 concentration system at the Rubisco are not very sensitive to high temperatures. The aforementioned reasons as above may explain the decrease in CO2assimilation when the temperature increases beyond that of the thermal optimum.
vii. Hardening afterward institute exposure to cool (≤ about 10°C) and high (≥ about 37°C) temperatures
Maintaining plants at cool or high temperatures causes, forth with the changes in photosynthesis described above, increase in their resistance to otherwise lethal temperatures(frost and high temperature). This is hardening.
In this procedure, temperature and light collaborate and the metabolic changes induced are sometimes very rapid (from minutes to hours).
Thus, cold hardening tin exist achieved at ordinary temperature by modulating the length of the light flow or its spectral composition in the red [16]. However, cold is all the same required to reach full hardening. Also the lack of low-cal in the cold prevents hardening to varying degrees.
At elevated temperatures : the transmitted signals actuate the synthesis of chaperone proteins (HSPs: Rut Schock Proteins) that repair denaturing proteins, also foreclose their coagulation or even help mark them for degradation.
At cool temperatures: the synthesis of chaperone proteins is also activated. It is accompanied by (i) the synthesis of "antifreeze" proteins that interfere with ice crystal formation and (ii) an increment in sugar synthesis tending to increase osmotic pressure in the cells.
Notation that the signaling pathways and their interactions inducing the genome response are merely partially known. The references given in "Learn More than" and an fastened Focus allow for further exploration of this evolving signal.
eight. Effects of temperature on photosynthesis: summary diagram
The summary diagram (Figure 15) classifies the effects of temperature on photosynthesis co-ordinate to the speed of temperature modify and the extent of its variation. Notation that hardening allows leaf maintenance in perennial leafage plants and therefore minimizes free energy loss under extreme temperature conditions.
Figure 15. Scheme classifying the effects of temperature on photosynthesis. [Source: Writer'southward diagram]
The rapidity of current climate 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 product to proceed the biosphere functioning.
nine. Messages to call up
The uptake of CO2 by a leaf has a thermal optimum shut to the average temperature of its growth environment.
This thermal optimum can alter rapidly when the conditions of the surround are durably modified: this is a procedure acclimatization.
This thermal optimum is on average less in C3 plants than in C4 plants: this is mainly due to photosynthetic fixation of atmospheric Otwo via Rubisco activity in C3 plants.
This optimum depends on the COii 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 leafage.
Subjected to cool or hot temperatures plants bring into play processeshardening to otherwise lethal temperatures. These processes involve poly peptide syntheses and changes in the fluidity of chloroplast and prison cell membranes.
Notes and references
Cover epitome. Sunset over the Sonora Arizona desert. [Source: royalty gratuitous / Pixabay]
[1] Meehl GA, Stocker TF, Collins WD, Riedlingstein P, Gaye AT, Gregory JM, Kitoh A, Knutti R, Spud JM, Noda A & Raper SCB (2007). Climate Change 2007: The Physical Science Footing. Contribution of Working Group I to the Fourth Cess Written report of the Intergovernmental Panel on Climatic change. Cambridge University Printing
[2] Yamori W, Hikosaka Grand & Way DA. (2014). Temperature response of photosynthesis in C3, C4, and CAM plants: temperature acclimation and temperature adaptation. Photosynthesis Res. 119, 101- 117.
[3] For example, when growing plants are subjected to drought, the amount of carbon they assimilate decreases initially considering leaf growth is inhibited. The mechanisms for CO2 fixation in the leaf are not then inhibited. Boyer JS (1970) Plant Physiol.46, 233-235
[4] The values of thermal optima given here, are from measurements fabricated in "normal air", containing 21% O2 and most 400 ppm CO2. When this is not the case the O2 and CO2 contents are shown. The CO2 uptake in air containing 21% Otwo is saturated from nearly 1200 ppm CO2 when calorie-free is close to saturation. The evaporative power of the air is also regulated in most cases during the measurements. It is estimated by the saturation deficit of the partial pressure of water vapor in the ambient air around the leaves.
[five] Plants from the same individual past vegetative reproduction. They are genetically identical.
[half-dozen] Ecotype: Plants of the same species from unlike environments, which, grown from seed to flower nether identical conditions bear witness different physiological characteristics.
[7] It fetches h2o from every bit far equally the water table, hence its name of phreatophyte establish.
[8] Pearcy RW (1971). Acclimation of photosynthetic and respiratory CO2 exchange to growth temperature in Atriplex lentiJormis (Torr.) Wats. Plant Physiol. 59, 795-799
[9] Yamasaki T, Yamakawa T, Yamane Y, koike H, Satoh One thousand & Katoh S. (2002) Temperature acclimation of photosynthesis and related changes in photosystem 2 electron transport in winter wheat. Found Physiol. 128 1087-1097.
[10] See notation #iv, section 2.2
[xi] Jordan DB & Ogren WL (1984). The CO2/Otwo specificity of ribulose 1,five-bisphosphate carboxylase/oxygenase. Dependence on ribulose bisphosphate concentration, pH and temperature. Planta161, 308-313
[12] Ehleringer JR, Sage RF, Flanagan LB & Pearcy RW (1991). Climatic change and the evolution of C4 photosynthesis. Trends in Ecology and Evolution 6, 95-99
[13] Cornic G & Massacci A (1996). Leafage photosynthesis nether drought stress. In Advances in Photosynthesis (vol 5) Photosynthesis and the surroundings, 347-366. Neil R Bakery (ed.) Kluwer Bookish publishers Dordrecht.
[xiv] Cornic G, Badeck F-Due west, Ghashghaie J & Manuel N (1999). Event of temperature on net COii 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 Thousand (eds) Crop development for cool and wet regions of Europe. European community. ISBN 92-828-6947-four.
[15] Crafts-Brandner SJ, van de Loo FJ & Salvucci ME (1997). The two forms of ribulose-1,5-bisphosphate carboxylase/oxygenase activase differ in sensitivity to elevated temperature. Constitute Physiol. 114, 439-444.
[16] Puhakainen T, Li C, Boije-Malm Grand, Kangasjärvi J, Heino P & Palva ET. (2004). Short-day potentiation of low temperature-induced cistron expression of a C-repeat-binding factor-controlled cistron during common cold acclimation in Silver Birch. Plant Physiol.136, 4299-4307
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To cite this article: CORNIC Gabriel (2022), Effects of temperature on photosynthesis, Encyclopedia of the Environment, [online ISSN 2555-0950] url : https://www.encyclopedie-environnement.org/en/life/effects-temperature-on-photosynthesis/.
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