Abstract
Upon hydration and dehydration, the vegetative tissue of Selaginella lepidophylla can reversibly swell and shrink to generate complex morphological transformations. Here, we investigate how structural and compositional properties at tissue and cell wall levels in S. lepidophylla lead to different stem curling profiles between inner and outer stems. Our results show that directional bending in both stem types is associated with cross-sectional gradients of tissue density, cell orientation and secondary cell wall composition between adaxial and abaxial stem sides. In inner stems, longitudinal gradients of cell wall thickness and composition affect tip-to-base tissue swelling and shrinking, allowing for more complex curling as compared to outer stems. Together, these features yield three-dimensional functional gradients that allow the plant to reproducibly deform in predetermined patterns that vary depending on the stem type. This study is the first to demonstrate functional gradients at different hierarchical levels combining to operate in a three-dimensional context.
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Claims 3
Claim · model-written summary
Directional bending in both stem types is associated with cross-sectional gradients in tissue density, cell orientation and secondary wall composition.
- Taxon
- Selaginella lepidophylla
Selaginella lepidophyllat0 - Organ / stage
- steme0
- Outcome
- directional bendinge0
- Qualifier
- cross-sectional gradients of tissue density, cell orientation and secondary cell wall compositione0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Our results show that directional bending in both stem types is associated with cross-sectional gradients of tissue density, cell orientation and secondary cell wall composition between adaxial and abaxial stem sides.”
t0“Selaginella lepidophylla”
Claim · model-written summary
In inner stems, longitudinal cell wall gradients affect swelling and shrinking and permit more complex curling than outer stems.
- Taxon
- Selaginella lepidophylla
Selaginella lepidophyllat0 - Organ / stage
- inner stemse0
- Outcome
- more complex curlinge0
- Qualifier
- longitudinal gradients of cell wall thickness and compositione0
- Comparator
- outer stemse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“In inner stems, longitudinal gradients of cell wall thickness and composition affect tip-to-base tissue swelling and shrinking, allowing for more complex curling as compared to outer stems.”
t0“Selaginella lepidophylla”
Claim · model-written summary
Three-dimensional functional gradients allow reproducible deformation patterns varying by stem type.
- Taxon
- Selaginella lepidophylla
Selaginella lepidophyllat0 - Outcome
- reproducibly deform in predetermined patternse0
- Qualifier
- three-dimensional functional gradientse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Together, these features yield three-dimensional functional gradients that allow the plant to reproducibly deform in predetermined patterns that vary depending on the stem type.”
t0“Selaginella lepidophylla”