PERSPECTIVE

Engineering photosynthesis, nature’s carbon

capture machine
Megan L. Matthews ID1,2*
1 Department of Civil and Environmental Engineering, University of Illinois at Urbana-Champaign, Urbana,
Illinois, United States of America, 2 Carl R Woese Institute for Genomic Biology, University of Illinois at
Urbana–Champaign, Urbana, Illinois, United States of America

* [email protected]

Mitigating climate change and sustainably feeding our growing pop-

ulation in the changing climate are 2 significant challenges facing the
global community. Engineering photosynthesis, nature’s carbon cap-
ture machinery, can help us surmount these threats.

For thousands of years, humans have sought to increase plant productivity to create better and
a1111111111 more abundant food. The Green Revolution in the second half of the 20th century saw us
a1111111111 make significant improvements in crop yields through increased use of fertilizers, irrigation,
a1111111111 and strategic breeding of crops to develop new high-yielding varieties. The impacts of this agri-
a1111111111 cultural revolution were vast and included averting hunger for millions of people [1]. Since
a1111111111 then, our population has continued to grow, passing 8 billion people in 2022, and is projected
to peak at around 10.4 billion people in 2086 [2]. Traditional approaches that have been used
to increase crop productivity are now plateauing. To feed this population, we need to develop
new approaches to further increase crop productivity, especially in regions of the world where
OPEN ACCESS people do not currently have reliable access to food.
Photosynthesis, the process plants use to convert light energy into chemical energy, is
Citation: Matthews ML (2023) Engineering
photosynthesis, nature’s carbon capture machine. responsible for nearly all life on Earth. Most terrestrial plants use the C3 photosynthetic path-
PLoS Biol 21(7): e3002183. https://doi.org/ way to harvest light and convert that solar radiation into chemical energy in the form of glu-
10.1371/journal.pbio.3002183 cose. Yet, photosynthesis is not an efficient process, and most terrestrial plants only convert
Academic Editor: Pamela C. Ronald, University of about 1% to 2% of the incoming light energy into chemical energy. As a comparison, commer-
California, Davis, UNITED STATES cially available solar panels convert around 20% of solar energy into electricity. For most of
history, there has not been a need for plants to improve on this 1% to 2% efficiency. However,
Published: July 11, 2023
if we want to further increase crop productivity, we need to increase the amount carbon that is
Copyright: © 2023 Megan L. Matthews. This is an being assimilated by the crops in the first place. Improving the efficiency of photosynthesis
open access article distributed under the terms of
would increase this source amount of assimilated carbon, which could then be transported to
the Creative Commons Attribution License, which
permits unrestricted use, distribution, and the yield or to a plant organ of interest.
reproduction in any medium, provided the original Beyond increasing crop production for food, improving photosynthetic efficiency would
author and source are credited. capture more CO2 from the atmosphere that can be stored in plant biomass and eventually
Funding: The author(s) received no specific sequestered in the soil or used to create bioproducts [3]. Removing CO2 from the atmosphere
funding for this work. is an important climate change mitigation strategy, and photosynthesis is a natural way of cap-
turing this carbon. To improve photosynthetic efficiency for greater carbon capture, we need
Competing interests: The authors have declared
that no competing interests exist. to be able to modify large modules or components of the photosynthesis machinery or even

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add new components all together [4]. These complex changes are difficult to achieve through
evolution or breeding but can, and have been, accomplished through plant engineering.
Energy is lost at multiple steps throughout photosynthesis, preventing plants from more
efficiently capturing carbon and converting solar energy to chemical energy. When it comes to
absorbing solar radiation, light-harvesting antenna in the chloroplasts only absorb radiation
that have wavelengths between 400 nm and 700 nm, which is roughly equivalent to the visible
light spectrum. The wavelengths in this spectrum account for less than 50% of the total energy
in solar radiation, and some of these wavelengths are reflected instead of absorbed, further
decreasing the amount of solar radiation that plants can convert to chemical energy. Moreover,
chloroplasts often experience fluctuating light throughout a day, transitioning through sun–
shade and shade–sun conditions due to clouds and other leaves or plants moving with the
wind. During these transitions, it can take several minutes for photosynthesis to return to max-
imal operation under the new light condition, which has been estimated to cost 10% to 40% of
potential CO2 assimilation [5].
Photorespiration and the stomatal and mesophyll conductances, which determine how
much CO2 reaches the chloroplasts, can also limit photosynthetic efficiency. Photorespiration
occurs when O2 instead of CO2 binds to Rubisco, a key enzyme in the carbon fixation pathway,
producing a different compound that must be recycled through the photorespiration pathway
to recover back some of the carbon for photosynthesis. This process can reduce photosynthetic
efficiency by 20% to 50% [6]. The amount of photorespiration that occurs is related to the ratio
of CO2 to O2 in the chloroplast, where higher ratios of CO2 to O2 result in less photorespira-
tion. This ratio is dependent on how easily CO2 can enter the leaf cells and chloroplasts, which
is determined by leaf anatomy and the stomatal and mesophyll conductances. Variations from
the C3 photosynthetic pathway have evolved independently in several plant species to avoid
photorespiration (Box 1).

Box 1. C3, C4, and CAM photosynthesis

Photosynthesis comes in several different forms, with C3, C4, and crassulacean acid
metabolism (CAM) pathways being the most common in plants. Approximately 85% of
plants, including major crops like soybean, rice, and wheat, use the C3 photosynthetic
pathway. In these plants, the photosynthetic reactions occur in the chloroplasts of leaf
mesophyll cells. CO2 enters the leaf through stomatal pores before making its way into
the chloroplasts. When these stomata open to take in CO2, water is also released from
the leaves through transpiration. As such, plants need to balance CO2 uptake with water
loss. Under drought conditions, plants will keep their stomata closed to prevent the loss
of water. This also prevents CO2 from entering the mesophyll cells, leading to an
increase in photorespiration, which occurs when Rubisco, a key enzyme in photosynthe-
sis, binds with O2 instead of CO2.
In response to photorespiration, the C4 and CAM pathways have evolved independently
in about 3% and 6% of flowering plant species, respectively [7]. C4 and CAM photosyn-
thetic pathways use more energy to convert solar energy to glucose than the C3 pathway,
but they have the benefit of concentrating CO2 around Rubisco, thereby increasing the
efficiency of the carboxylation reaction as CO2 is no longer competing with O2 to bind
to Rubisco. In C4 plants, like maize and sugarcane, CO2 enters the mesophyll cells where
it is converted to a carbon intermediate that is transported from the mesophyll cells into
another leaf cell, the bundle sheath. In the bundle sheath cells, the carbon intermediate is
then converted back into CO2 and binds to Rubisco and the carbon fixation pathway

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proceeds. In CAM plants, like pineapple and aloe vera, the stomata only open during the
night to prevent water loss. The CO2 that enters the mesophyll chloroplasts is then con-
verted into an intermediate. During the day, the carbon intermediates are converted
back to CO2, where they then bind with Rubisco, and, as with the C4 process, carbon fix-
ation proceeds. Since C4 and CAM plants store CO2 as intermediates, they have more
flexibility in when to open their stomata, allowing them to conserve water better than C3
plants. C4 and CAM photosynthetic pathways tend to be found in plants that are native
to hot, arid environments where water loss can be a major problem.

The current state of photosynthesis (in)efficiency, combined with the number of avenues
for its improvement, underscores the potential for engineering photosynthesis to increase car-
bon capture for food and storage. Scientists have already demonstrated in greenhouse and
field experiments how we can increase photosynthesis through engineering different parts of
the process, including increasing how fast photosynthetic machinery responds to fluctuating
light [8,9], adding shorter and less energetically costly photorespiration recovery pathways [6],
and manipulating photosynthetic enzymes to increase reaction rates [10]. In addition to these
modifications, there are still a vast number of options for photosynthesis engineering that are
being explored, from expanding the range of wavelengths that can be absorbed by the light-
harvesting antenna [11] to introducing carboxysomes, carbon-concentrating compartments
from blue-green algae, into C3 photosynthetic pathways [12].
Given the complexity of photosynthesis and the many different avenues towards engineer-
ing it for increased efficiency and carbon capture, mathematical models have been useful tools
for identifying which strategies are promising for implementation [9]. Models will continue to
be important resources to identify and evaluate different engineering strategies for increasing
carbon capture at the field scale, across developmental stages, and under future climate scenar-
ios. Models can also be used to investigate the predicted combined impact from stacking mul-
tiple engineered traits, such as increased light absorption and increased enzymatic reaction
rates or less costly photorespiration.
Despite successful greenhouse and field experiments demonstrating that we can engineer
plant photosynthesis to increase carbon assimilation, there remain gaps to translating these
and future engineering strategies from research labs and fields into crops that can be planted
by farmers around the world. For photosynthesis engineering to have a role in mitigating cli-
mate change and sustainably feeding our growing population, we need to bridge these regula-
tory, economic, social, and political gaps through specific translational grants, industry and
foundation partnerships, and initiatives designed to accelerate the pipeline from discovery sci-
ence to societal impact. We are living in a period of significant change and upheaval caused by
our own actions and inactions. To address these challenges and meet the needs of the next cen-
tury, we must innovate and implement beyond our traditional tools for increasing crop pro-
ductivity. Engineering photosynthesis for improved efficiency and increased carbon capture,
when used in conjunction with other scientific and societal advances, can help us solve at least
2 significant global challenges of the next century.

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