# Convergent evolution increases boron transport through SNPs and tandem duplications at BOR1 and BOR2 in Arabidopsis thaliana

**Authors:** Emmanuel Tergemina, Célia Neto, Md Mamunur Rashid, Herculano Dinis, David E. Salt, Angela M. Hancock

PMC · DOI: 10.1073/pnas.2525676123 · Proceedings of the National Academy of Sciences of the United States of America · 2026-03-23

## TL;DR

Plants from boron-poor volcanic islands evolved similar ways to boost boron uptake, offering clues for developing resilient crops.

## Contribution

Discovery of convergent evolution in BOR1 and BOR2 genes through SNPs and duplications in Arabidopsis thaliana.

## Key findings

- Multiple populations evolved increased leaf boron via mutations in BOR1 and BOR2.
- Tandem duplications at BOR1 arose independently in different populations.
- Combining GWAS and recombinant populations improves detection of genetic variants in natural settings.

## Abstract

Boron is an essential nutrient for plant growth, but many soils lack sufficient amounts, limiting crop production worldwide. To understand how plants adapt to boron-deficient environments, we studied wild mustard plants that naturally colonized volcanic islands, where boron is typically scarce. We identified a case of convergent evolution, wherein different plant populations independently evolved to increase boron concentrations in their leaves through distinct genetic changes in genes controlling boron transport. This finding reveals a key biological mechanism that may help plants survive in boron-poor soils, providing valuable insights for developing more resilient crops that can thrive in challenging agricultural environments.

Boron (B) is a crucial micronutrient, particularly in volcanic soils where its deficiency hampers agriculture. Here, we investigate the genetic basis of leaf B accumulation in natural populations of Arabidopsis thaliana that colonized volcanic islands in Cape Verde. Using a combination of genome-wide association studies (GWAS) and mapping in a recombinant intercross population, we identified a case of convergent phenotypic evolution in which multiple variants in the two principal B transporter genes, BOR1 and BOR2, increase leaf B accumulation in parallel. These include multiple tandem duplications at BOR1 that arose independently in different populations. Overall, this study reveals a remarkable case of convergent evolution occurring within a relatively short time scale, where different types of de novo mutations at B efflux transporter genes achieve similar phenotypic outcomes. Further, our findings show that integrating recombinant populations with GWAS in natural populations can improve power to overcome allelic heterogeneity.

## Linked entities

- **Genes:** BOR1 (HCO3- transporter family) [NCBI Gene 819329], SIX5 (SIX homeobox 5) [NCBI Gene 147912]
- **Chemicals:** boron (PubChem CID 5462311)
- **Species:** Arabidopsis thaliana (taxon 3702)

## Full-text entities

- **Genes:** NRAMP1 (natural resistance-associated macrophage protein 1) [NCBI Gene 844422] {aka ATNRAMP1, F23A5.18, F23A5_18, PMIT1, natural resistance-associated macrophage protein 1}, NIP5;1 (NOD26-like intrinsic protein 5;1) [NCBI Gene 826630] {aka AtNIP5;1, F7L13.6, NLM6, NLM8, NOD26-LIKE MIP 6, NOD26-LIKE MIP 8}, RMR1 (receptor homology region transmembrane domain ring H2 motif protein 1) [NCBI Gene 836748] {aka ARABIDOPSIS THALIANA RECEPTOR HOMOLOGY REGION TRANSMEMBRANE DOMAIN RING H2 MOTIF PROTEIN 1, ATRMR1, JR700, K2A18.24, K2A18_24, REMEMBR-H2 PROTEIN JR700}, BOR4 (HCO3- transporter family) [NCBI Gene 838116] {aka ARABIDOPSIS THALIANA REQUIRES HIGH BORON 4, ATBOR4, F9L1.41, F9L1_41, REQUIRES HIGH BORON 4}, BOR1 (HCO3- transporter family) [NCBI Gene 819329] {aka AtBOR1, REQUIRES HIGH BORON 1, T3D7.3}, AT3G62270 (HCO3- transporter family) [NCBI Gene 825400] {aka BOR2, REQUIRES HIGH BORON 2}, PP2A (serine/threonine protein phosphatase 2A) [NCBI Gene 843333] {aka F20P5.30, F20P5_30, TYPE 2A SERINE/THREONINE PROTEIN PHOSPHATASE, serine/threonine protein phosphatase 2A}
- **Diseases:** toxicity (MESH:D064420), tissue necrosis (MESH:D009336), TD (MESH:D058674), growth defects (MESH:D006130), sterility (MESH:D007246), CVI (MESH:C537835), B (MESH:D006509)
- **Chemicals:** Ti (MESH:D014025), TRIzol (MESH:C411644), water (MESH:D014867), B (MESH:D001895), lipid (MESH:D008055), Cr (MESH:D002857), borate (MESH:D001881), hydrogen (MESH:D006859), AM1906 (-), RG-II (MESH:C042492), ethanol (MESH:D000431), PNAS (MESH:D020135), polystyrene (MESH:D011137), FAM (MESH:C031179), Boric acid (MESH:C032688)
- **Species:** Brassica napus (oilseed rape, species) [taxon 3708], Zygnema sp. 'M' (species) [taxon 2494501], Hepatovirus A (no rank) [taxon 12092], Zea mays (maize, species) [taxon 4577], Arabidopsis thaliana (mouse-ear cress, species) [taxon 3702], Helianthus annuus (common sunflower, species) [taxon 4232]
- **Mutations:** S313, S313G, 313G

## Full text

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## Figures

6 figures with captions in the complete paper: https://tomesphere.com/paper/PMC13037888/full.md

## References

80 references — full list in the complete paper: https://tomesphere.com/paper/PMC13037888/full.md

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Source: https://tomesphere.com/paper/PMC13037888