Abstract
Biological shapes are often produced by the iterative generation of repeated units. The mechanistic basis of such iteration is an area of intense investigation. Leaf development in the model plant Arabidopsis is one such example where the repeated generation of leaf margin protrusions, termed serrations, is a key feature of final shape. However, the regulatory logic underlying this process is unclear. Here, we use a combination of developmental genetics and computational modeling to show that serration development is the morphological read-out of a spatially distributed regulatory mechanism, which creates interspersed activity peaks of the growth-promoting hormone auxin and the cup-shaped cotyledon2 (CUC2) transcription factor. This mechanism operates at the growing leaf margin via a regulatory module consisting of two feedback loops working in concert. The first loop relates the transport of auxin to its own distribution, via polar membrane localization of the pinformed1 (PIN1) efflux transporter. This loop captures the potential of auxin to generate self-organizing patterns in diverse developmental contexts. In the second loop, CUC2 promotes the generation of PIN1-dependent auxin activity maxima while auxin represses CUC2 expression. This CUC2-dependent loop regulates activity of the conserved auxin efflux module in leaf margins to generate stable serration patterns. Conceptualizing leaf margin development via this mechanism also helps to explain how other developmental regulators influence leaf shape.
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Claims 6
Claim · model-written summary
Serration development is attributed to spatial regulatory peaks of auxin and CUC2 activity.
- Taxon
- Arabidopsis thaliana
Arabidopsis thalianat0 - Gene
- cup-shaped cotyledon2 (CUC2)e0
- Compound
- auxine0
- Outcome
- serration developmente0
- Qualifier
- spatially distributed regulatory mechanisme0
Not reportedOrgan / stage
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“serration development is the morphological read-out of a spatially distributed regulatory mechanism, which creates interspersed activity peaks of the growth-promoting hormone auxin and the cup-shaped cotyledon2 (CUC2) transcription factor.”
t0“Arabidopsis thaliana”
Claim · model-written summary
The proposed mechanism has two feedback loops operating at the growing leaf margin.
- Taxon
- Arabidopsis thaliana
Arabidopsis thalianat0 - Organ / stage
- growing leaf margine0
- Outcome
- two feedback loops working in concerte0
- Qualifier
- regulatory modulee0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“This mechanism operates at the growing leaf margin via a regulatory module consisting of two feedback loops working in concert.”
t0“Arabidopsis thaliana”
Claim · model-written summary
PIN1 polar membrane localization links auxin transport to its distribution.
- Taxon
- Arabidopsis thaliana
Arabidopsis thalianat0 - Gene
- pinformed1 (PIN1)e0
- Compound
- auxine0
- Outcome
- transport of auxin to its own distributione0
- Qualifier
- polar membrane localizatione0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“The first loop relates the transport of auxin to its own distribution, via polar membrane localization of the pinformed1 (PIN1) efflux transporter.”
t0“Arabidopsis thaliana”
Claim · model-written summary
CUC2 promotes PIN1-dependent auxin activity maxima.
- Taxon
- Arabidopsis thaliana
Arabidopsis thalianat0 - Gene
- CUC2e0
- Compound
- auxine0
- Outcome
- PIN1-dependent auxin activity maximae0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“CUC2 promotes the generation of PIN1-dependent auxin activity maxima”
t0“Arabidopsis thaliana”
Claim · model-written summary
Auxin represses CUC2 expression.
- Taxon
- Arabidopsis thaliana
Arabidopsis thalianat0 - Gene
- CUC2e0
- Compound
- auxine0
- Outcome
- CUC2 expressione0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“auxin represses CUC2 expression”
t0“Arabidopsis thaliana”
Claim · model-written summary
The CUC2-dependent loop regulates an auxin efflux module to generate stable serration patterns.
- Taxon
- Arabidopsis thaliana
Arabidopsis thalianat0 - Organ / stage
- leaf marginse0
- Gene
- CUC2e0
- Outcome
- stable serration patternse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“This CUC2-dependent loop regulates activity of the conserved auxin efflux module in leaf margins to generate stable serration patterns.”
t0“Arabidopsis thaliana”