---
OA_place: publisher
OA_type: free access
_id: '22570'
abstract:
- lang: eng
  text: Atmospheric carbon dioxide concentration ([CO 2 ]) is increasing, which increases
    leaf-scalephotosynthesis and intrinsic water-use efﬁciency. These direct responses
    have the potential toincrease plant growth, vegetation biomass, and soil organic
    matter; transferring carbon from theatmosphere into terrestrial ecosystems (a
    carbon sink). A substantial global terrestrial carbon sinkwould slow the rate
    of [CO 2] increase and thus climate change. However, ecosystem CO2responses are
    complex or confounded by concurrent changes in multiple agents of global changeand
    evidence for a [CO 2]-driven terrestrial carbon sink can appear contradictory.
    Here wesynthesize theory and broad, multidisciplinary evidence for the effects
    of increasing [CO 2](iCO 2) on the global terrestrial carbon sink. Evidence suggests
    a substantial increase in globalphotosynthesis since pre-industrial times. Established
    theory, supported by experiments,indicates that iCO 2 is likely responsible for
    about half of the increase. Global carbon budgeting,atmospheric data, and forest
    inventories indicate a historical carbon sink, and these apparentiCO 2 responses
    are high in comparison to experiments and predictions from theory. Plantmortality
    and soil carbon iCO 2 responses are highly uncertain. In conclusion, a range of
    evidencesupports a positive terrestrial carbon sink in response to iCO2 , albeit
    with uncertain magnitudeand strong suggestion of a role for additional agents
    of global change.
article_processing_charge: No
article_type: original
author:
- first_name: Anthony P.
  full_name: Walker, Anthony P.
  last_name: Walker
- first_name: Martin G.
  full_name: De Kauwe, Martin G.
  last_name: De Kauwe
- first_name: Ana
  full_name: Bastos, Ana
  last_name: Bastos
- first_name: Soumaya
  full_name: Belmecheri, Soumaya
  last_name: Belmecheri
- first_name: Katerina
  full_name: Georgiou, Katerina
  last_name: Georgiou
- first_name: Ralph F.
  full_name: Keeling, Ralph F.
  last_name: Keeling
- first_name: Sean M.
  full_name: McMahon, Sean M.
  last_name: McMahon
- first_name: Belinda E.
  full_name: Medlyn, Belinda E.
  last_name: Medlyn
- first_name: David J. P.
  full_name: Moore, David J. P.
  last_name: Moore
- first_name: Richard J.
  full_name: Norby, Richard J.
  last_name: Norby
- first_name: Sönke
  full_name: Zaehle, Sönke
  last_name: Zaehle
- first_name: Kristina J.
  full_name: Anderson‐Teixeira, Kristina J.
  last_name: Anderson‐Teixeira
- first_name: Giovanna
  full_name: Battipaglia, Giovanna
  last_name: Battipaglia
- first_name: Roel J. W.
  full_name: Brienen, Roel J. W.
  last_name: Brienen
- first_name: Kristine G.
  full_name: Cabugao, Kristine G.
  last_name: Cabugao
- first_name: Maxime
  full_name: Cailleret, Maxime
  last_name: Cailleret
- first_name: Elliott
  full_name: Campbell, Elliott
  last_name: Campbell
- first_name: Josep G.
  full_name: Canadell, Josep G.
  last_name: Canadell
- first_name: Philippe
  full_name: Ciais, Philippe
  last_name: Ciais
- first_name: Matthew E.
  full_name: Craig, Matthew E.
  last_name: Craig
- first_name: David S.
  full_name: Ellsworth, David S.
  last_name: Ellsworth
- first_name: Graham D.
  full_name: Farquhar, Graham D.
  last_name: Farquhar
- first_name: Simone
  full_name: Fatichi, Simone
  id: cf8e546b-a9b0-11f0-a43b-aa89ed1b56d6
  last_name: Fatichi
- first_name: Joshua B.
  full_name: Fisher, Joshua B.
  last_name: Fisher
- first_name: David C.
  full_name: Frank, David C.
  last_name: Frank
- first_name: Heather
  full_name: Graven, Heather
  last_name: Graven
- first_name: Lianhong
  full_name: Gu, Lianhong
  last_name: Gu
- first_name: Vanessa
  full_name: Haverd, Vanessa
  last_name: Haverd
- first_name: Kelly
  full_name: Heilman, Kelly
  last_name: Heilman
- first_name: Martin
  full_name: Heimann, Martin
  last_name: Heimann
- first_name: Bruce A.
  full_name: Hungate, Bruce A.
  last_name: Hungate
- first_name: Colleen M.
  full_name: Iversen, Colleen M.
  last_name: Iversen
- first_name: Fortunat
  full_name: Joos, Fortunat
  last_name: Joos
- first_name: Mingkai
  full_name: Jiang, Mingkai
  last_name: Jiang
- first_name: Trevor F.
  full_name: Keenan, Trevor F.
  last_name: Keenan
- first_name: Jürgen
  full_name: Knauer, Jürgen
  last_name: Knauer
- first_name: Christian
  full_name: Körner, Christian
  last_name: Körner
- first_name: Victor O.
  full_name: Leshyk, Victor O.
  last_name: Leshyk
- first_name: Sebastian
  full_name: Leuzinger, Sebastian
  last_name: Leuzinger
- first_name: Yao
  full_name: Liu, Yao
  last_name: Liu
- first_name: Natasha
  full_name: MacBean, Natasha
  last_name: MacBean
- first_name: Yadvinder
  full_name: Malhi, Yadvinder
  last_name: Malhi
- first_name: Tim R.
  full_name: McVicar, Tim R.
  last_name: McVicar
- first_name: Josep
  full_name: Penuelas, Josep
  last_name: Penuelas
- first_name: Julia
  full_name: Pongratz, Julia
  last_name: Pongratz
- first_name: A. Shafer
  full_name: Powell, A. Shafer
  last_name: Powell
- first_name: Terhi
  full_name: Riutta, Terhi
  last_name: Riutta
- first_name: Manon E. B.
  full_name: Sabot, Manon E. B.
  last_name: Sabot
- first_name: Juergen
  full_name: Schleucher, Juergen
  last_name: Schleucher
- first_name: Stephen
  full_name: Sitch, Stephen
  last_name: Sitch
- first_name: William K.
  full_name: Smith, William K.
  last_name: Smith
- first_name: Benjamin
  full_name: Sulman, Benjamin
  last_name: Sulman
- first_name: Benton
  full_name: Taylor, Benton
  last_name: Taylor
- first_name: César
  full_name: Terrer, César
  last_name: Terrer
- first_name: Margaret S.
  full_name: Torn, Margaret S.
  last_name: Torn
- first_name: Kathleen K.
  full_name: Treseder, Kathleen K.
  last_name: Treseder
- first_name: Anna T.
  full_name: Trugman, Anna T.
  last_name: Trugman
- first_name: Susan E.
  full_name: Trumbore, Susan E.
  last_name: Trumbore
- first_name: Phillip J.
  full_name: van Mantgem, Phillip J.
  last_name: van Mantgem
- first_name: Steve L.
  full_name: Voelker, Steve L.
  last_name: Voelker
- first_name: Mary E.
  full_name: Whelan, Mary E.
  last_name: Whelan
- first_name: Pieter A.
  full_name: Zuidema, Pieter A.
  last_name: Zuidema
citation:
  ama: Walker AP, De Kauwe MG, Bastos A, et al. Integrating the evidence for a terrestrial
    carbon sink caused by increasing atmospheric CO2. <i>New Phytologist</i>. 2021;229(5):2413-2445.
    doi:<a href="https://doi.org/10.1111/nph.16866">10.1111/nph.16866</a>
  apa: Walker, A. P., De Kauwe, M. G., Bastos, A., Belmecheri, S., Georgiou, K., Keeling,
    R. F., … Zuidema, P. A. (2021). Integrating the evidence for a terrestrial carbon
    sink caused by increasing atmospheric CO2. <i>New Phytologist</i>. Wiley. <a href="https://doi.org/10.1111/nph.16866">https://doi.org/10.1111/nph.16866</a>
  chicago: Walker, Anthony P., Martin G. De Kauwe, Ana Bastos, Soumaya Belmecheri,
    Katerina Georgiou, Ralph F. Keeling, Sean M. McMahon, et al. “Integrating the
    Evidence for a Terrestrial Carbon Sink Caused by Increasing Atmospheric CO2.”
    <i>New Phytologist</i>. Wiley, 2021. <a href="https://doi.org/10.1111/nph.16866">https://doi.org/10.1111/nph.16866</a>.
  ieee: A. P. Walker <i>et al.</i>, “Integrating the evidence for a terrestrial carbon
    sink caused by increasing atmospheric CO2,” <i>New Phytologist</i>, vol. 229,
    no. 5. Wiley, pp. 2413–2445, 2021.
  ista: Walker AP, De Kauwe MG, Bastos A, Belmecheri S, Georgiou K, Keeling RF, McMahon
    SM, Medlyn BE, Moore DJP, Norby RJ, Zaehle S, Anderson‐Teixeira KJ, Battipaglia
    G, Brienen RJW, Cabugao KG, Cailleret M, Campbell E, Canadell JG, Ciais P, Craig
    ME, Ellsworth DS, Farquhar GD, Fatichi S, Fisher JB, Frank DC, Graven H, Gu L,
    Haverd V, Heilman K, Heimann M, Hungate BA, Iversen CM, Joos F, Jiang M, Keenan
    TF, Knauer J, Körner C, Leshyk VO, Leuzinger S, Liu Y, MacBean N, Malhi Y, McVicar
    TR, Penuelas J, Pongratz J, Powell AS, Riutta T, Sabot MEB, Schleucher J, Sitch
    S, Smith WK, Sulman B, Taylor B, Terrer C, Torn MS, Treseder KK, Trugman AT, Trumbore
    SE, van Mantgem PJ, Voelker SL, Whelan ME, Zuidema PA. 2021. Integrating the evidence
    for a terrestrial carbon sink caused by increasing atmospheric CO2. New Phytologist.
    229(5), 2413–2445.
  mla: Walker, Anthony P., et al. “Integrating the Evidence for a Terrestrial Carbon
    Sink Caused by Increasing Atmospheric CO2.” <i>New Phytologist</i>, vol. 229,
    no. 5, Wiley, 2021, pp. 2413–45, doi:<a href="https://doi.org/10.1111/nph.16866">10.1111/nph.16866</a>.
  short: A.P. Walker, M.G. De Kauwe, A. Bastos, S. Belmecheri, K. Georgiou, R.F. Keeling,
    S.M. McMahon, B.E. Medlyn, D.J.P. Moore, R.J. Norby, S. Zaehle, K.J. Anderson‐Teixeira,
    G. Battipaglia, R.J.W. Brienen, K.G. Cabugao, M. Cailleret, E. Campbell, J.G.
    Canadell, P. Ciais, M.E. Craig, D.S. Ellsworth, G.D. Farquhar, S. Fatichi, J.B.
    Fisher, D.C. Frank, H. Graven, L. Gu, V. Haverd, K. Heilman, M. Heimann, B.A.
    Hungate, C.M. Iversen, F. Joos, M. Jiang, T.F. Keenan, J. Knauer, C. Körner, V.O.
    Leshyk, S. Leuzinger, Y. Liu, N. MacBean, Y. Malhi, T.R. McVicar, J. Penuelas,
    J. Pongratz, A.S. Powell, T. Riutta, M.E.B. Sabot, J. Schleucher, S. Sitch, W.K.
    Smith, B. Sulman, B. Taylor, C. Terrer, M.S. Torn, K.K. Treseder, A.T. Trugman,
    S.E. Trumbore, P.J. van Mantgem, S.L. Voelker, M.E. Whelan, P.A. Zuidema, New
    Phytologist 229 (2021) 2413–2445.
das_tickbox: '1'
date_created: 2026-07-27T12:30:24Z
date_published: 2021-03-01T00:00:00Z
date_updated: 2026-08-06T08:39:39Z
day: '01'
doi: 10.1111/nph.16866
extern: '1'
external_id:
  pmid:
  - '32789857'
intvolume: '       229'
issue: '5'
keyword:
- Beta factor
- Carbon dioxide
- CO2 fertilization
- CO2-fertilization hypothesis
- Free-air CO2 enrichment (FACE)
- Global carbon cycle
- Land–atmosphere feedback
- Terrestrial ecosystems
language:
- iso: eng
main_file_link:
- open_access: '1'
  url: https://doi.org/10.1111/nph.16866
month: '03'
oa: 1
oa_version: Published Version
page: 2413-2445
pmid: 1
publication: New Phytologist
publication_identifier:
  eissn:
  - 1469-8137
  issn:
  - 0028-646X
publication_status: published
publisher: Wiley
quality_controlled: '1'
scopus_import: '1'
status: public
title: Integrating the evidence for a terrestrial carbon sink caused by increasing
  atmospheric CO2
type: journal_article
user_id: ba8df636-2132-11f1-aed0-ed93e2281fdd
volume: 229
year: '2021'
...
---
OA_place: publisher
OA_type: free access
_id: '22536'
abstract:
- lang: eng
  text: Increasing concentrations of atmospheric carbon dioxide are expected to affect
    carbon assimilation and evapotranspiration (ET), ultimately driving changes in
    plant growth, hydrology, and the global carbon balance. Direct leaf biochemical
    effects have been widely investigated, whereas indirect effects, although documented,
    elude explicit quantification in experiments. Here, we used a mechanistic model
    to investigate the relative contributions of direct (through carbon assimilation)
    and indirect (via soil moisture savings due to stomatal closure, and changes in
    leaf area index) effects of elevated CO2 across a variety of ecosystems. We specifically
    determined which ecosystems and climatic conditions maximize the indirect effects
    of elevated CO2. The simulations suggest that the indirect effects of elevated
    CO2 on net primary productivity are large and variable, ranging from less than
    10% to more than 100% of the size of direct effects. For ET, indirect effects
    were, on average, 65% of the size of direct effects. Indirect effects tended to
    be considerably larger in water-limited ecosystems. As a consequence, the total
    CO2 effect had a significant, inverse relationship with the wetness index and
    was directly related to vapor pressure deficit. These results have major implications
    for our understanding of the CO2 response of ecosystems and for global projections
    of CO2 fertilization, because, although direct effects are typically understood
    and easily reproducible in models, simulations of indirect effects are far more
    challenging and difficult to constrain. Our findings also provide an explanation
    for the discrepancies between experiments in the total CO2 effect on net primary
    productivity.
article_processing_charge: No
article_type: original
author:
- first_name: Simone
  full_name: Fatichi, Simone
  id: cf8e546b-a9b0-11f0-a43b-aa89ed1b56d6
  last_name: Fatichi
- first_name: Sebastian
  full_name: Leuzinger, Sebastian
  last_name: Leuzinger
- first_name: Athanasios
  full_name: Paschalis, Athanasios
  last_name: Paschalis
- first_name: J. Adam
  full_name: Langley, J. Adam
  last_name: Langley
- first_name: Alicia
  full_name: Donnellan Barraclough, Alicia
  last_name: Donnellan Barraclough
- first_name: Mark J.
  full_name: Hovenden, Mark J.
  last_name: Hovenden
citation:
  ama: Fatichi S, Leuzinger S, Paschalis A, Langley JA, Donnellan Barraclough A, Hovenden
    MJ. Partitioning direct and indirect effects reveals the response of water-limited
    ecosystems to elevated CO. <i>Proceedings of the National Academy of Sciences</i>.
    2016;113(45):12757-12762. doi:<a href="https://doi.org/10.1073/pnas.1605036113">10.1073/pnas.1605036113</a>
  apa: Fatichi, S., Leuzinger, S., Paschalis, A., Langley, J. A., Donnellan Barraclough,
    A., &#38; Hovenden, M. J. (2016). Partitioning direct and indirect effects reveals
    the response of water-limited ecosystems to elevated CO. <i>Proceedings of the
    National Academy of Sciences</i>. National Academy of Sciences. <a href="https://doi.org/10.1073/pnas.1605036113">https://doi.org/10.1073/pnas.1605036113</a>
  chicago: Fatichi, Simone, Sebastian Leuzinger, Athanasios Paschalis, J. Adam Langley,
    Alicia Donnellan Barraclough, and Mark J. Hovenden. “Partitioning Direct and Indirect
    Effects Reveals the Response of Water-Limited Ecosystems to Elevated CO.” <i>Proceedings
    of the National Academy of Sciences</i>. National Academy of Sciences, 2016. <a
    href="https://doi.org/10.1073/pnas.1605036113">https://doi.org/10.1073/pnas.1605036113</a>.
  ieee: S. Fatichi, S. Leuzinger, A. Paschalis, J. A. Langley, A. Donnellan Barraclough,
    and M. J. Hovenden, “Partitioning direct and indirect effects reveals the response
    of water-limited ecosystems to elevated CO,” <i>Proceedings of the National Academy
    of Sciences</i>, vol. 113, no. 45. National Academy of Sciences, pp. 12757–12762,
    2016.
  ista: Fatichi S, Leuzinger S, Paschalis A, Langley JA, Donnellan Barraclough A,
    Hovenden MJ. 2016. Partitioning direct and indirect effects reveals the response
    of water-limited ecosystems to elevated CO. Proceedings of the National Academy
    of Sciences. 113(45), 12757–12762.
  mla: Fatichi, Simone, et al. “Partitioning Direct and Indirect Effects Reveals the
    Response of Water-Limited Ecosystems to Elevated CO.” <i>Proceedings of the National
    Academy of Sciences</i>, vol. 113, no. 45, National Academy of Sciences, 2016,
    pp. 12757–62, doi:<a href="https://doi.org/10.1073/pnas.1605036113">10.1073/pnas.1605036113</a>.
  short: S. Fatichi, S. Leuzinger, A. Paschalis, J.A. Langley, A. Donnellan Barraclough,
    M.J. Hovenden, Proceedings of the National Academy of Sciences 113 (2016) 12757–12762.
das_tickbox: '1'
date_created: 2026-07-27T12:30:24Z
date_published: 2016-11-08T00:00:00Z
date_updated: 2026-08-06T08:08:46Z
day: '08'
doi: 10.1073/pnas.1605036113
extern: '1'
external_id:
  pmid:
  - '27791074'
intvolume: '       113'
issue: '45'
keyword:
- carbon dioxide
- modeling
- FACE
- soil moisture
- evapotranspiration
language:
- iso: eng
main_file_link:
- open_access: '1'
  url: https://doi.org/10.1073/pnas.1605036113
month: '11'
oa: 1
oa_version: Published Version
page: 12757-12762
pmid: 1
publication: Proceedings of the National Academy of Sciences
publication_identifier:
  eissn:
  - 1091-6490
  issn:
  - 0027-8424
publication_status: published
publisher: National Academy of Sciences
quality_controlled: '1'
scopus_import: '1'
status: public
title: Partitioning direct and indirect effects reveals the response of water-limited
  ecosystems to elevated CO
type: journal_article
user_id: ba8df636-2132-11f1-aed0-ed93e2281fdd
volume: 113
year: '2016'
...
