Terrestrial plants are regularly subjected to strong temperature variations. These variations can reach 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 assimilation of CO
2
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 slow 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.
The current increase in greenhouse gas emissions will cause an increase in atmospheric temperature of 2 to 3°C in the next 50 years (see
A carbon cycle disrupted by human activities
). At the same time, heat waves and extreme heat periods will be more frequent and of longer duration
[1]
. Agricultural production and the functioning of forests will therefore be greatly affected. Models based on large-scale observations indicate that, in the absence of agronomic adaptation, the decrease in crop yields can reach 17% for each 1°C increase in the temperature of the growing season
[2]
.
The production of higher plants
depends in particular (but not only
[3]
) on
leaf photosynthesis
(see
Shedding light on photosynthesis
& The
The path of carbon in photosynthesis
). CO
2
enters the leaf where its reduction in the chloroplasts is accompanied by O
2
production. Its entry is almost exclusively through the stomata (Figure 1). For each molecule of CO
2
absorbed, 50 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
Leaf transpiration and heat protection
).
Figure 1. During photosynthesis, CO2 is absorbed and O2 is released mainly through the stomatal opening (ostiole). The water 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: Author’s diagram]
The leaf is a
converter of solar energy into chemical energy
and, like any energy converter, requires a permanent
cooling system
.
The climate changes that are currently occurring make it necessary to understand the effects of temperature on photosynthesis.
2. The thermal optimum of photosynthesis
2.1. Diagram of the thermal response
Photosynthetic CO
2
uptake varies with temperature. In most cases its response to temperature is
rapidly reversible between
about
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 10°C and above 34°C plants start to set up
protective mechanisms
. For these extreme values, CO
2
assimilation is often unstable and can be cancelled more or less quickly: the leaf is then under stress (Figure 2).
2.2. A thermal optimum based on the average temperature of the environment
Plants in 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 high temperatures.
Figure 3.
Deschampsia antarctica
is one 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 CO
2
assimilation
[4]
in
Deschampsia antarctica
(Figure 3) and
Colobanthus quitensis
, the only two Antarctic flowering plants, is between 8 and 15°C, while it is around 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.
2.3. Acclimatization to the thermal conditions of the environment
Figure 4. Variations in CO2 assimilation as a function of leaf temperature, in a plant 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 also found in individuals of the same species growing at different temperatures. Figure 4 shows CO
2
assimilation in pea grown at 10 or 25°C.
In the first case (cultivation at 10°C) the thermal optimum is about 16°C, while it is higher than 25°C in the second (cultivation at 25°C). At low temperatures, CO
2
assimilation is higher in plants grown at 10°C.
In this case the adjustment to cool conditions is a gain for the plant.
2.4. Acclimatization can be rapid
Figure 5. Remth
(Hammada scoparia
), a characteristic plant of the Wadi Rum desert (Jordan). [Source: Ji-Elle, CC BY-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
In general, these changes can be measured in both growing and mature leaves, with the response being of
greater amplitude
in
growing
leaves.
2.5. Heat-sensitive
versus
cold-sensitive species
Warm acclimation of cool-adapted species (or ecotypes
[6]
) occurs with an
increase in thermal optimum
but a general
decrease
in
photosynthesis
.
This is, for example, the case of
Atriplex sabulosa
. One can then wonder about the interest of this change. The opposite may be true for plants strictly adapted to warm conditions, such as
Tridestomia oblongifolia
. Figure 7 illustrates the case of
Atripex lentiformis
,
[7]
a perennial leafy plant, which occurs in California in both Death 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 night (in red in Figure 7).
Figure 7. Variations in CO2 assimilation of Atriplex lentiformis ecotypes from a warm environment (A) and a cool, moist environment (B). [Source: Author’s diagram, after Pearcy (1977)] Right, Atripex lentiformis (salt bush) [Source: Forest & Kim Starr, CC BY 3.0, via Wikimedia Commons]
Under the alternating 43°C day and 30°C night (blue in Figure 7), only the desert ecotype shows plasticity, maintaining high CO
2
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]
.
C4 plants,
of which there are traces only from the end 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
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 20 ° C, the
photosynthesis
of C4 plants is
on average lower
than that of C3 plants.
3. CO
2
assimilation results from the interaction of processes whose response to temperature is different
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 diffusion of CO
2
from the ambient air to the chloroplasts: its speed increases with temperature.
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 made 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 sugar with 5 carbon atoms (Ribulose 1,5-bisphosphate: C5). Benson-Calvin cycle: allows the regeneration of C5, and at the same time 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 two origins: (1) oxidation of water in the lumen by 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 process) which uses reducing power (in the form 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
2
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
CO
2
assimilation when the latter is
saturating
[9]
.
The activity of PSII
has a thermal optimum identical to that of the electron transfer chain.
PSI activity is not inhibited at high temperatures
(above 30°C, up to 45°C) where it remains stable or even increases: it 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 complex that allows the oxidation of water (see Figure 9).
The thermal response of electron transfer
is
similar
in
C3 and C4
plants. However, there are organizational differences between these two types of plants (see
The path of carbon in photosynthesis
).
The supply of energy cannot therefore explain 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 O
2
on CO
2
assimilation of C3 and C4 plants
In normal air
[10]
, 21% O
2
(+ N
2
) + 360 ppm CO
2
: the thermal optimum is 27°C in Maize (C4 plant), while it is only 22°C in Pea (C3 plant) (Figure 10): the
thermal optimum
of the
C4
plant is
higher
than that of the C3 plant (see also section 2.6).
In an oxygen-deficient atmosphere
, 1% O
2
(+ N
2
) + 360 ppm CO
2
: the CO
2
uptake of Maize is not affected, while that of Pea is stimulated above about 17°C, with a shift in its thermal optimum to near that of Maize.
In
C3
plants
, atmospheric oxygen inhibits CO
2
uptake when the leaf temperature is sufficiently high,
whereas it has no effect
(or negligible effect) in C3 plants
Figure 10. Variation of CO2 assimilation measured in leaves of Pea (A; Pisum sativum) and Maize (B; Zea mays) as a function of leaf temperature. The plants were grown in natural light at a temperature of 20 ± 2°C. [Source: Author’s diagram – royalty-free image / Pixabay]
Note that the variation in electron transfer estimated
in vivo
, by measuring chlorophyll fluorescence emission as a function of temperature, is very similar in 1% and 21% O
2
in Pea:
the variation in thermal optimum
is therefore
not due to a change in photochemistry
.
4.3. Rubisco properties explain the difference in response
Case of C3 plants
CO
2
and O
2
compete to occupy the active sites of Rubisco: This enzyme has a carboxylase function and an oxygenase function. CO
2
enters the Benson-Calvin cycle and the
photosynthetic fixation of O
2
is at the origin of a metabolic pathway responsible for
photorespiration
(Figure 11; see also
The path of carbon in photosynthesis
).
CO
2
occupies a high number of active sites on the Rubisco when the O
2
content of the ambient air is low (1% for example) or that of CO
2
is high.
O
2
is mainly fixed if its content increases
or if that of CO
2
decreases (the latter then releases active sites which are then occupied by O
2
)
.
In
normal air
, there are two reasons why
O
2
fixation increases
(and consequently CO
2
fixation decreases) when the
temperature increases
[11]
.
The affinity of Rubisco for
CO2
decreases more than that for O
2
; a factor that favours the assimilation of O
2
.
The water solubility coefficient of CO
2
decreases more than that of O
2
, leading to a more rapid decrease in the amount of CO
2
than O
2
in the chloroplast; this is a factor that favours O
2
fixation.
In an
O
2
-poor atmosphere
(Figure 10),
competition between O
2
and CO
2
is very reduced. Energy is then used mainly for CO
2
assimilation, which increases in value until around 30°C and then decreases as the energy supply decreases (see section 4.1).
In
normal air,
the effect of O
2
on photosynthetic CO
2
fixation (Figure 11) is very low (or even 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 1,5-bisphosphate) in a C3 plant. APG: 3-phosphoglyceric acid, 3 C compound; TP: Trioses phosphate. The carbon leaves the Calvin cycle to feed the synthesis of sucrose. PG: phosphoglycolate, 2C compound. Two 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 O
2
which then consumes an increasing part of the energy produced by the activity of the photosystems. This energy is therefore no longer available for CO
2
fixation, which reaches its maximum value around 22°C.
Case of C4 plants
.
CO
2
is 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
current atmospheric content
.
Under these conditions, photosynthetic O
2
fixation is weak or even non-existent because the active sites of the Rubisco are all occupied by CO
2
.
The energy
supplied by the activity of the photosystems is therefore
used only in the fixation of CO
2
when the
leaf temperature increases,
explaining the higher thermal optimum in this type of plant.
C4 plants evolved
from C3 plants
during the global
decrease
in
atmospheric CO
2
content at the end of the Tertiary 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 establishment of a CO
2
concentration mechanism is an
advantage
because 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 environmental parameters
5.1. The CO
2
content in the atmosphere
The thermal optimum increases with increasing ambient CO
2
content. In the case shown in Figure 12, it increases from about 10°C when the content is 100 ppm to more than 30°C when it is 800 ppm.
Figure 13. Human activities lead to an increase 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 activity of the vegetation. [Source : Royalty-free image / Pixabay]
In a world with steadily
increasing
atmospheric CO
2
(Figure 13), the thermal optimum of C3 plants is expected to increase. This does not mean, however, that plant production will then be higher (see note 3 section 1):
episodes of high heat
will, like droughts, certainly be
more frequent
.
5.2. Lack of water
The photosynthetic apparatus is resistant to drought
. It retains all its capacity to absorb CO
2
on the Rubisco, and to produce energy until the leaves have lost about 30% of their water
[13]
.
CO
2
uptake decreases in this range of water loss, because the
stomata close
(see Focus L
eaf transpiration and heat protection
). This closure
slows down the entry of CO
2
into the leaf and consequently leads to a
decrease of the CO
2
content in the mesophyll.
However, the O
2
content in the chloroplasts remains high. Indeed, its content in the atmosphere (21% or 210,000 ppm) is, compared to that of CO
2
(@ 400 ppm), very high and in any case sufficient for a very substantial quantity to pass through the epidermis even when the stomata are closed.
The competition between CO
2
and O
2
for the occupation of the active sites of the Rubisco is thus in
favour of O
2
.
Figure 14. A, Variations in CO2 assimilation as a function 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 rate estimated on the same leaves by measuring the chlorophyll fluorescence emission. [Source: Author’s diagram, after Cornic et al. ref. 14]
Therefore, the thermal optimum for photosynthesis must lower in C3 plants that dry out.
This is shown in Figure 14A, in which the thermal optimum drops from about 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 by water loss in the range shown (Figure 14B). When water loss is 20%, 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, CO
2
assimilation 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 activity of some enzymes controlling this turnover, notably that of a Fructose 1,6-bisphosphate (see Figures 9 and 11).
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
(see section 4.1), resulting in reduced energy production (reducing power 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 CO
2
accumulation 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 CO
2
. However, other factors are at play since CO
2
assimilation measured (1) in an atmosphere with little or no photorespiration (ambient O
2
content of 1%), and (2) measured in a normal atmosphere in a C4 plant decreases in both cases (Figure 10).
Several reasons can be given:
The slowing down of PSII activity
leading to that of the electron transfer chain from PSII to PSI.
To perform its role
Rubisco must be 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 appear in some plants subjected to periods of high 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-1,4-bisphosphate), which is structurally close to RuBP (see Figures 9 and 11), are synthesized.
In C4 plants
(case of Maize) the activation and activity of enzymes that participate in the CO
2
concentration system at the Rubisco are not very sensitive to high temperatures. The same reasons as above may explain the decrease in CO
2
assimilation when the temperature increases beyond that of the thermal optimum.
7. Hardening after plant exposure to cool (≤ about 10°C) and high (≥ about 37°C) temperatures
Maintaining plants at cool or high temperatures causes, along with the changes in photosynthesis described above,
increase in their resistance to otherwise lethal temperatures
(frost and high temperature). This is
hardening.
In this process, temperature and light interact and the metabolic changes induced are sometimes very rapid (from minutes to hours).
Thus, cold hardening can be achieved at ordinary temperature by
modulating the length of the light period or its spectral composition in the red
[16]
. However, cold is still required to achieve full hardening.
Also the lack of light 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 prevent 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 increase in sugar synthesis tending to increase osmotic pressure in the cells.
Note that the signaling pathways and their interactions inducing the genome response are
only partially
known. The references given in “Learn More” and an attached Focus allow for further exploration of this evolving point.
8. Effects of temperature on photosynthesis: summary diagram
The summary diagram (Figure 15) classifies the effects of temperature on photosynthesis according to the speed of temperature change and the extent of its variation. Note that hardening allows leaf maintenance in perennial leaf plants and therefore minimizes energy loss under extreme temperature conditions.
Figure 15. Scheme classifying the effects of temperature on photosynthesis. [Source: Author’s 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 production to keep the biosphere functioning.
9.
Messages to remember
The uptake of CO
2
by a leaf has a
thermal optimum
close to the average temperature of its growth environment.
This thermal optimum can
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 O
2
via Rubisco activity in C3 plants.
This optimum depends on the
CO
2
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 cool 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.
Notes and references
Cover image.
Sunset over the Sonora Arizona desert. [Source: royalty free / Pixabay]
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and CO
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[5]
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[6]
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), contractually linked to the University of Grenoble Alpes and Grenoble INP, and sponsored by the French Academy of Sciences.
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Author(s)
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, Honorary Professor, Laboratoire d’écophysiologie des plantes, Université Paris-Saclay
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How is the carbon from carbon dioxide – CO2 – present in the atmosphere integrated…
Jean-François MOROT-GAUDRY
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Global climate change is a major concern. It is based on the scientific community’s statement…
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, General engineer of bridges, waters and forests, former climatologist researcher at Météo-France, CNRM (Centre National de Recherches Météorologiques).
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图7. 左图为生活在温暖环境（A）和凉爽湿润环境（B）的两种大滨藜（Atriplex lentiformis）生态型二氧化碳吸收速率的差异。[图片来源：作者自制，见Pearcy（1977）]。右图为大滨藜（盐木）[图片来源：Forest & Kim Starr, CC BY 3.0, 通过维基共享资源]
在白天43 °C、夜间30 °C的生长环境中（图7中蓝色部分），只有沙漠生态型表现出适应性，在新生长条件下保持了高的二氧化碳吸收速率；而滨海生态型在所有叶片温度下的光合速率都较低，只有最适温度的升高体现了它一定的适应能力
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Author(s)
CORNIC Gabriel
, Honorary Professor, Laboratoire d’écophysiologie des plantes, Université Paris-Saclay
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How is the carbon from carbon dioxide – CO2 – present in the atmosphere integrated…
Jean-François MOROT-GAUDRY
, INRA Emeritus Research Director, the Jean-Pierre Bourgin Institute, INRA Versailles, Member of the Académie d’Agriculture.
Global climate change is a major concern. It is based on the scientific community’s statement…
Serge PLANTON
, General engineer of bridges, waters and forests, former climatologist researcher at Météo-France, CNRM (Centre National de Recherches Météorologiques).
