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PHOTOSYNTHESIS
Just the right temperature
A global analysis of the relationship between photosynthesis and temperature identifies key similarities and
differences when scaling from leaves to ecosystems and suggests that carbon uptake by vegetation may be able to
adjust to a warming world.
Danielle A. Way
P
hotosynthesis is a key player in the
global carbon cycle, with terrestrial
vegetation absorbing ~120 gigatons
of carbon from the atmosphere annually1.
A critical question is, therefore, how will
climate warming impact this enormous
flux? If warming stimulates photosynthesis,
this could slow climate change by pulling
carbon dioxide out of the atmosphere,
helping to offset anthropogenic emissions.
But if increasing temperatures suppress
global photosynthesis, this could reduce
plant productivity and increase the rate
of climate change. Addressing this issue
at large spatial scales is difficult because,
although we know how warmer conditions
alter the thermal sensitivity of carbon
dioxide uptake at the leaf level2,3, it is unclear
whether these findings can be extrapolated
to entire ecosystems. In this issue of Nature
Ecology & Evolution, Huang et al.4 provide
the first global analysis of how ecosystem
photosynthesis relates to temperature,
uncovering tantalizing parallels with
findings from the leaf level.
As one scales up in biological complexity,
translating knowledge from one level to
the next becomes increasingly difficult.
The biochemical processes underlying leaf
photosynthesis peak at 35–40 °C (ref. 5),
but neither leaf- nor ecosystem-level
carbon dioxide uptake is purely driven by
photosynthetic enzyme kinetics (Fig. 1a).
Carbon dioxide must diffuse through
stomata to reach the photosynthetic
machinery, and stomata close under hot,
dry conditions, reducing carbon dioxide
availability at high temperatures and thus
creating physiological constraints on
productivity. Additionally, carbon dioxide
is emitted from leaves via respiration and
photorespiration. As such, the thermal
optimum of leaf net photosynthesis (which
accounts for carbon dioxide uptake via
photosynthetic biochemistry, stomatal
limitations on carbon dioxide supply, and
respiratory and photorespiratory carbon
dioxide losses) is 25–30 °C for most
plants6, well below the thermal optimum
for photosynthetic biochemistry. These
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and other constraints are likely to also
affect thermal optima when scaled up to
ecosystem photosynthesis, but by
how much?
Using eddy covariance (which measures
ecosystem gas exchange) and remote
sensing (which quantifies spectral data
that correlate with canopy photosynthetic
activity), Huang et al. found that, globally,
the optimum temperature for ecosystem
photosynthesis was 23 °C — close to
that of leaf net photosynthesis. Although
leaf net photosynthesis and ecosystem
photosynthesis are not exactly comparable
(as the former includes leaf respiration and
the latter does not; see Fig. 1a), the similarity
between their thermal optima emphasizes
that realized photosynthetic rates in
individual leaves and ecosystems are both
constrained by stomatal conductance, leaf
age and other factors at high temperatures.
Huang et al. also examined whether
the temperature at which ecosystem
photosynthesis peaked was correlated with
local climate conditions, which would
indicate that carbon uptake in vegetation
was tuned to the prevailing thermal
environment. In leaves, the photosynthetic
thermal optimum acclimates to prevailing
temperatures: for every 1 °C increase
in growth temperature, the optimum
temperature for net photosynthesis increases
by 0.55–0.62 °C (refs. 2,7) (Fig. 1b,c). In
other words, leaves adjust photosynthesis
to achieve maximum rates at temperatures
similar to those experienced in their
environment. This acclimation is largely
driven by changes in the temperature
sensitivity of the enzymatic processes in
photosynthesis, whose thermal optima
rise by 0.63–0.71 °C per 1 °C in growth
temperature7 (Fig. 1c). Huang et al.4
demonstrate that ecosystem photosynthesis
also peaks at higher temperatures at warmer
sites, at a rate remarkably similar to that
found in leaves: 0.61 °C per 1 °C increase
in air temperature. Although the data in
Huang et al. cannot tell us to what degree
this global pattern is related to adaptation
to local thermal regimes versus thermal
plasticity of photosynthesis, acclimation
plays a larger role in dictating the optimum
temperature of leaf net photosynthesis than
does evolution7. If ecosystem carbon dioxide
uptake follows a similar pattern, then as the
climate warms, the temperature optimum
of ecosystem photosynthesis should also
increase, thereby mitigating some of the
potential negative impacts of climate change
on the carbon cycle.
The news that ecosystem-level carbon
uptake is likely to adjust to warmer
temperatures is heartening, but the finding
that shifts in the thermal optimum do not
keep pace with warming is worrisome. In
tropical regions, Huang et al. found that
the optimum temperature for ecosystem
photosynthesis was already near the
maximum air temperature experienced
during the growing season (Fig. 1b).
Since the thermal optimum of ecosystem
photosynthesis rises by only 0.61 °C per
1 °C increase in air temperature, any
warming at these latitudes should reduce
carbon uptake. In contrast, sites at higher
latitudes had thermal optima for ecosystem
photosynthesis that were well above the
maximum growing season temperatures4,
providing plants in these regions with a
large thermal buffer against the impacts
of warming on carbon uptake. However,
climate warming is predicted to be more
extreme at high latitudes, with warming
of 8 °C or more possible by the end of the
century8. High-latitude vegetation may
therefore be at just as much of a risk for
suppressed photosynthesis over the next few
decades as tropical ecosystems, since rapid
climate warming would erase the thermal
buffer currently benefiting these species.
The work by Huang et al. is an important
step toward understanding how warming
will affect the drawdown of carbon dioxide
by the biosphere. Their data emphasize
how even small temperature increases
could reduce carbon dioxide uptake in
the tropics and imply that rapid warming
at high latitudes poses a similar danger to
these regions. Perhaps most importantly,
the finding that warming generates
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a b
Ecosystem respiration 4 °C

CO2 out
Photosynthesis
2.4 °C
CO2 in

or ecosystem photosynthesis
Leaf net photosynthesis
Ecosystem
photosynthesis

Photorespiration
CO2 out

Air temperature (°C)

c
Biochemical demand
(0.71 °C per 1 °C)

Change in thermal optimum (°C)

Ecosystem photosynthesis
(0.61 °C per 1 °C)
Leaf net photosynthesis
(0.55 °C per 1 °C)
1:1

Photosynthesis Photorespiration Respiration

CO2 in CO2 out CO2 out

Leaf net photosynthesis

Change in air temperature (°C)

Fig. 1 | Influence of temperature on leaf- and ecosystem-level photosynthesis. a, A comparison of the carbon dioxide fluxes involved in leaf net
photosynthesis and ecosystem photosynthesis (as measured with eddy covariance). At the leaf level, net photosynthesis includes: (1) carbon dioxide
diffusion through stomata; (2) uptake of carbon dioxide by enzymes in photosynthesis; (3) the release of carbon dioxide by photorespiration; and (4) the
release of carbon dioxide by respiration. Ecosystem photosynthesis accounts for the first three of these processes, whereas leaf respiration is combined with
other autotrophic and heterotrophic respiration fluxes to derive ecosystem respiration. b, Leaf net photosynthesis and ecosystem photosynthesis are both
sensitive to changing temperatures. The blue line shows how carbon dioxide uptake responds to changes in air temperature for vegetation that grows at cool
temperatures (the maximum growing season temperature is indicated by the blue arrow), with maximum rates at a thermal optimum (Topt, blue circle). Under
warmer temperatures (the maximum growing season temperature is indicated by the red arrow), the thermal performance curve of photosynthesis shifts
(shown by the red line), increasing Topt (red circle). But since Topt increases by ~0.6 °C per 1 °C of warming, a 4 °C increase in air temperature leads to only a
2.4 °C increase in Topt. The difference between the air temperature and Topt therefore shrinks under warming, reducing the thermal buffer for carbon dioxide
uptake (delineated by the blue and red solid bars). c, The Topt values for photosynthetic biochemical demand for carbon dioxide, leaf net photosynthesis and
ecosystem photosynthesis all show similar shifts as temperatures increase, but, since these shifts do not keep pace with warming, there is an increasing
mismatch between the thermal environment and the optimal temperature for carbon dioxide uptake.

an increasing mismatch between air
temperatures and the thermal optimum
of photosynthesis, a mismatch that is of a
similar magnitude across multiple biological
scales, emphasizes the pressing need to
determine why photosynthetic acclimation
cannot keep pace with warming. ❐ https://doi.org/10.1038/s41559-019-0877-3
Danielle A. Way1,2,3
1
Department of Biology, University of Western
Ontario, London, Ontario, Canada. 2Nicholas School
of the Environment, Duke University, Durham, NC,
USA. 3Division of Plant Sciences, Research School of 4. Huang, M. et al. Nat. Ecol. Evol. https://doi.org/10.1038/s41559-
019-0838-x (2019).
Biology, Australian National University, Canberra, 5. Kattge, J. & Knorr, W. Plant Cell Environ. 30,
Australian Capital Territory, Australia. 1176–1190 (2007).
e-mail: dway4@uwo.ca 6. Sage, R. F. & Kubien, D. S. Plant Cell Environ. 30,
1086–1106 (2007).
7. Kumarathunge, D. et al. New Phytol. 222, 768–784 (2019).
Published: xx xx xxxx 8. Ciais, P. et al. 2013. Carbon and other biogeochemical cycles. in
Climate Change 2013: The Physical Science Basis. Contribution
of Working Group I to the Fifth Assessment Report of the
Intergovernmental Panel on Climate Change (eds Tignor, M. et al.)
References Ch. 6. (Cambridge University Press, 2013).
1. Le Quéré, C. et al. Earth Syst. Sci. Data 8, 605–649 (2016).
2. Yamori, W., Hikosaka, K. & Way, D. A. Photosynth. Res. 119,
101–117 (2014). Competing interests
3. Way, D. A. & Yamori, W. Photosynth. Res. 119, 89–100 (2014). The author declares no competing interests.

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