Abstract
Abstract Atmospheric carbon dioxide ( CO 2 ) directly determines the rate of plant photosynthesis and indirectly effects plant productivity and fitness and may therefore act as a selective pressure driving evolution, but evidence to support this contention is sparse. Using Plantago lanceolata L. seed collected from a naturally high CO 2 spring and adjacent ambient CO 2 control site, we investigated multigenerational response to future, elevated atmospheric CO 2 . Plants were grown in either ambient or elevated CO 2 (700 μmol mol −1 ), enabling for the first time, characterization of the functional and population genomics of plant acclimation and adaptation to elevated CO 2 . This revealed that spring and control plants differed significantly in phenotypic plasticity for traits underpinning fitness including above‐ground biomass, leaf size, epidermal cell size and number and stomatal density and index. Gene expression responses to elevated CO 2 (acclimation) were modest [33–131 genes differentially expressed ( DE )], whilst those between control and spring plants (adaptation) were considerably larger (689–853 DE genes). In contrast, population genomic analysis showed that genetic differentiation between spring and control plants was close to zero, with no fixed differences, suggesting that plants are adapted to their native CO 2 environment at the level of gene expression. An unusual phenotype of increased stomatal index in spring but not control plants in elevated CO 2 correlated with altered expression of stomatal patterning genes between spring and control plants for three loci ( YODA , CDKB 1;1 and SCRM 2 ) and between ambient and elevated CO 2 for four loci ( ER , YODA , MYB 88 and BCA 1 ). We propose that the two positive regulators of stomatal number ( SCRM 2 ) and CDKB 1;1 when upregulated act as key controllers of stomatal adaptation to elevated CO 2 . Combined with significant transcriptome reprogramming of photosynthetic and dark respiration and enhanced growth in spring plants, we have identified the potential basis of plant adaptation to high CO 2 likely to occur over coming decades.
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Claims 12
Claim · model-written summary
Spring and control Plantago plants differed significantly in phenotypic plasticity for above-ground biomass, leaf size and epidermal cell size.
- Taxon
- Plantago lanceolata L.
Plantago lanceolatae0 - Material
- spring and control plantse2
- Outcome
- phenotypic plasticitye2
- Qualifier
- above‐ground biomass, leaf size, epidermal cell sizee2
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Using Plantago lanceolata L. seed collected from a naturally high CO 2 spring and adjacent ambient CO 2 control site”
e2“spring and control plants differed significantly in phenotypic plasticity for traits underpinning fitness including above‐ground biomass, leaf size, epidermal cell size and number and stomatal density and index”
Claim · model-written summary
Spring and control Plantago plants differed significantly in phenotypic plasticity for epidermal cell number and stomatal density and index.
- Taxon
- Plantago lanceolata L.
Plantago lanceolatae0 - Material
- spring and control plantse2
- Outcome
- phenotypic plasticitye2
- Qualifier
- stomatal density and indexe2
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Using Plantago lanceolata L. seed collected from a naturally high CO 2 spring and adjacent ambient CO 2 control site”
e2“spring and control plants differed significantly in phenotypic plasticity for traits underpinning fitness including above‐ground biomass, leaf size, epidermal cell size and number and stomatal density and index”
Claim · model-written summary
Gene expression responses to elevated CO2 (acclimation) were modest, with 33-131 differentially expressed genes.
- Taxon
- Plantago lanceolata L.
Plantago lanceolatae0 - Treatment
- elevated CO 2e1
- Outcome
- gene expression responses to elevated CO 2 (acclimation)e1
- Value
- 33–131 genes differentially expressede1
- Qualifier
- modeste1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Using Plantago lanceolata L. seed collected from a naturally high CO 2 spring and adjacent ambient CO 2 control site”
e1“Gene expression responses to elevated CO 2 (acclimation) were modest [33–131 genes differentially expressed ( DE )], whilst those between control and spring plants (adaptation) were considerably larger (689–853 DE genes)”
Claim · model-written summary
Gene expression differences between control and spring plants (adaptation) were considerably larger, with 689-853 DE genes.
- Taxon
- Plantago lanceolata L.
Plantago lanceolatae0 - Material
- control and spring plantse1
- Outcome
- gene expression responsese1
- Value
- 689–853 DE genese1
- Qualifier
- considerably largere1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Using Plantago lanceolata L. seed collected from a naturally high CO 2 spring and adjacent ambient CO 2 control site”
e1“Gene expression responses to elevated CO 2 (acclimation) were modest [33–131 genes differentially expressed ( DE )], whilst those between control and spring plants (adaptation) were considerably larger (689–853 DE genes)”
Claim · model-written summary
Genetic differentiation between spring and control Plantago plants was close to zero, with no fixed differences.
- Taxon
- Plantago lanceolata L.
Plantago lanceolatae0 - Material
- spring and control plantse3
- Outcome
- genetic differentiation between spring and control plantse3
- Qualifier
- close to zero, with no fixed differencese3
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Using Plantago lanceolata L. seed collected from a naturally high CO 2 spring and adjacent ambient CO 2 control site”
e3“population genomic analysis showed that genetic differentiation between spring and control plants was close to zero, with no fixed differences”
Claim · model-written summary
Plantago plants are adapted to their native CO2 environment at the level of gene expression.
- Taxon
- Plantago lanceolata L.
Plantago lanceolatae0 - Outcome
- adapted to their native CO 2 environmente7
- Qualifier
- at the level of gene expressione7
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Using Plantago lanceolata L. seed collected from a naturally high CO 2 spring and adjacent ambient CO 2 control site”
e7“suggesting that plants are adapted to their native CO 2 environment at the level of gene expression”
Claim · model-written summary
Stomatal index increased in spring plants but not control plants under elevated CO2.
- Taxon
- Plantago lanceolata L.
Plantago lanceolatae0 - Material
- springe4
- Treatment
- elevated CO 2e4
- Outcome
- stomatal indexe4
- Direction
- increasede4
- Comparator
- control plantse4
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Using Plantago lanceolata L. seed collected from a naturally high CO 2 spring and adjacent ambient CO 2 control site”
e4“An unusual phenotype of increased stomatal index in spring but not control plants in elevated CO 2 correlated with altered expression of stomatal patterning genes between spring and control plants for three loci ( YODA , CDKB 1;1 and SCRM 2 ) and between ambient and elevated CO 2 for four loci ( ER , YODA , MYB 88 and BCA 1 )”
Claim · model-written summary
Stomatal patterning gene expression was altered between spring and control plants at three loci (YODA, CDKB1;1 and SCRM2).
- Taxon
- Plantago lanceolata L.
Plantago lanceolatae0 - Material
- spring and control plantse4
- Gene
- YODA , CDKB 1;1 and SCRM 2e4
- Outcome
- expression of stomatal patterning genese4
- Direction
- alterede4
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Using Plantago lanceolata L. seed collected from a naturally high CO 2 spring and adjacent ambient CO 2 control site”
e4“An unusual phenotype of increased stomatal index in spring but not control plants in elevated CO 2 correlated with altered expression of stomatal patterning genes between spring and control plants for three loci ( YODA , CDKB 1;1 and SCRM 2 ) and between ambient and elevated CO 2 for four loci ( ER , YODA , MYB 88 and BCA 1 )”
Claim · model-written summary
Stomatal patterning gene expression was altered between ambient and elevated CO2 at four loci (ER, YODA, MYB88 and BCA1).
- Taxon
- Plantago lanceolata L.
Plantago lanceolatae0 - Treatment
- elevated CO 2e4
- Gene
- ER , YODA , MYB 88 and BCA 1e4
- Outcome
- expression of stomatal patterning genese4
- Direction
- alterede4
- Comparator
- ambiente4
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Using Plantago lanceolata L. seed collected from a naturally high CO 2 spring and adjacent ambient CO 2 control site”
e4“An unusual phenotype of increased stomatal index in spring but not control plants in elevated CO 2 correlated with altered expression of stomatal patterning genes between spring and control plants for three loci ( YODA , CDKB 1;1 and SCRM 2 ) and between ambient and elevated CO 2 for four loci ( ER , YODA , MYB 88 and BCA 1 )”
Claim · model-written summary
SCRM2 and CDKB1;1, when upregulated, may be key controllers of stomatal adaptation to elevated CO2.
- Taxon
- Plantago lanceolata L.
Plantago lanceolatae0 - Gene
- SCRM 2e5
- Outcome
- stomatal adaptation to elevated CO 2e5
- Qualifier
- when upregulatede5
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Using Plantago lanceolata L. seed collected from a naturally high CO 2 spring and adjacent ambient CO 2 control site”
e5“We propose that the two positive regulators of stomatal number ( SCRM 2 ) and CDKB 1;1 when upregulated act as key controllers of stomatal adaptation to elevated CO 2”
Claim · model-written summary
Spring plants showed significant transcriptome reprogramming of photosynthesis and dark respiration.
- Taxon
- Plantago lanceolata L.
Plantago lanceolatae0 - Material
- spring plantse6
- Outcome
- transcriptome reprogramming of photosynthetic and dark respiratione6
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Using Plantago lanceolata L. seed collected from a naturally high CO 2 spring and adjacent ambient CO 2 control site”
e6“Combined with significant transcriptome reprogramming of photosynthetic and dark respiration and enhanced growth in spring plants”
Claim · model-written summary
Spring plants showed enhanced growth.
- Taxon
- Plantago lanceolata L.
Plantago lanceolatae0 - Material
- spring plantse6
- Outcome
- growthe6
- Direction
- enhancede6
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Using Plantago lanceolata L. seed collected from a naturally high CO 2 spring and adjacent ambient CO 2 control site”
e6“Combined with significant transcriptome reprogramming of photosynthetic and dark respiration and enhanced growth in spring plants”