Abstract
As a physical response to water loss during drought, inner Selaginella lepidophylla stems curl into a spiral shape to prevent photoirradiation damage to their photosynthetic surfaces. Curling is reversible and involves hierarchical deformation, making S. lepidophylla an attractive model with which to study water-responsive actuation. Investigation at the organ and tissue level has led to the understanding that the direction and extent of stem curling can be partially attributed to stiffness gradients between adaxial and abaxial stem sides at the nanoscale. Here, we examine cell wall elasticity to understand how it contributes to the overall stem curling. We compare the measured elastic moduli along the stem length and between adaxial and abaxial stem sides using atomic force microscopy nano-indentation testing. We show that changes in cortex secondary cell wall development lead to cell wall stiffness gradients from stem tip to base, and also between adaxial and abaxial stem sides. Changes in cortical cell wall morphology and secondary cell wall composition are suggested to contribute to the observed stiffness gradients.
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Claims 4
Claim · model-written summary
Cell wall stiffness gradients occur in Selaginella lepidophylla stems from stem tip to base.
- Taxon
- Selaginella lepidophylla
Selaginella lepidophyllat0 - Organ / stage
- steme0
- Outcome
- cell wall stiffness gradients from stem tip to basee0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“resurrection plant Selaginella lepidophylla”
e0“We show that changes in cortex secondary cell wall development lead to cell wall stiffness gradients from stem tip to base, and also between adaxial and abaxial stem sides”
Claim · model-written summary
Changes in cortex secondary cell wall development lead to cell wall stiffness gradients in Selaginella lepidophylla stems.
- Taxon
- Selaginella lepidophylla
Selaginella lepidophyllat0 - Outcome
- cell wall stiffness gradientse0
- Qualifier
- changes in cortex secondary cell wall developmente0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“resurrection plant Selaginella lepidophylla”
e0“We show that changes in cortex secondary cell wall development lead to cell wall stiffness gradients from stem tip to base, and also between adaxial and abaxial stem sides”
Claim · model-written summary
Cell wall stiffness gradients occur between adaxial and abaxial stem sides of Selaginella lepidophylla.
- Taxon
- Selaginella lepidophylla
Selaginella lepidophyllat0 - Organ / stage
- adaxial and abaxial stem sidese0
- Outcome
- cell wall stiffness gradientse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“resurrection plant Selaginella lepidophylla”
e0“We show that changes in cortex secondary cell wall development lead to cell wall stiffness gradients from stem tip to base, and also between adaxial and abaxial stem sides”
Claim · model-written summary
Changes in cortical cell wall morphology and secondary cell wall composition may contribute to the observed stiffness gradients.
- Taxon
- Selaginella lepidophylla
Selaginella lepidophyllat0 - Outcome
- stiffness gradientse1
- Qualifier
- cortical cell wall morphology and secondary cell wall compositione1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“resurrection plant Selaginella lepidophylla”
e1“Changes in cortical cell wall morphology and secondary cell wall composition are suggested to contribute to the observed stiffness gradients”