Calcium dominance, ion regulation, and metabolic defenses underlie salt tolerance in the halophyte Azima sarmentosa


Abstract

Background: Soil salinity is a major stressor limiting plant productivity, yet halophytes have evolved diverse mechanisms to cope with excess salt. Azima sarmentosa, a woody halophyte native to Southeast Asia, thrives in saline, calcium-poor soils, but its tolerance mechanisms remain poorly understood. This study examined anatomical, physiological, and biochemical traits of A. sarmentosa across a natural salinity gradient.
Methods: Soils, stems, mature leaves, and young leaves were analyzed using ion quantification, SEM–EDS/EDX, Synchrotron radiation X-ray tomographic microscopy (SRXTM), FT-IR spectroscopy, and multivariate analyses (correlation, PCA) to evaluate Ca, Na, and metabolic traits associated with stress tolerance.
Results: Despite low soil Ca 2+ , plants maintained high Ca 2+ /Na + ratios and produced abundant Ca-oxalate (CaOx) crystals in leaves and stems, indicating selective uptake and biomineralization. SEM–EDS/EDX confirmed Ca-rich deposits and salt glands on both leaf surfaces, while SRXTM showed their three-dimensional distribution within tissues. Young leaves accumulated high levels of proline, phenolics, and flavonoids, reinforcing osmotic adjustment and antioxidant protection. FT-IR spectra confirmed phenolic groups. Correlation and PCA revealed antagonism: Ca-associated traits (pigments, proline, flavonoids) opposed Na/Cl and acetone-derived phenolics, highlighting divergence between Ca-driven protection and Na-linked stress.
Conclusion: A. sarmentosa withstands salinity through an integrative, calcium-centered strategy involving selective Ca 2+ uptake, Ca-oxalate biomineralization, salt secretion, and metabolic defenses. Unlike halophytes that rely mainly on sodium sequestration, this species exhibits a distinctive Ca-based adaptation. Ca-oxalate formation not only immobilizes Ca 2+ but also sequesters CO2-derived oxalate, linking ionic regulation with carbon cycling and extending the ecological significance of calcium-centered tolerance.
Ask to review this manuscript

Notes for potential reviewers

  • Volunteering is not a guarantee that you will be asked to review. There are many reasons: reviewers must be qualified, there should be no conflicts of interest, a minimum of two reviewers have already accepted an invitation, etc.
  • This is NOT OPEN peer review. The review is single-blind, and all recommendations are sent privately to the Academic Editor handling the manuscript. All reviews are published and reviewers can choose to sign their reviews.
  • What happens after volunteering? It may be a few days before you receive an invitation to review with further instructions. You will need to accept the invitation to then become an official referee for the manuscript. If you do not receive an invitation it is for one of many possible reasons as noted above.

  • PeerJ does not judge submissions based on subjective measures such as novelty, impact or degree of advance. Effectively, reviewers are asked to comment on whether or not the submission is scientifically and technically sound and therefore deserves to join the scientific literature. Our Peer Review criteria can be found on the "Editorial Criteria" page - reviewers are specifically asked to comment on 3 broad areas: "Basic Reporting", "Experimental Design" and "Validity of the Findings".
  • Reviewers are expected to comment in a timely, professional, and constructive manner.
  • Until the article is published, reviewers must regard all information relating to the submission as strictly confidential.
  • When submitting a review, reviewers are given the option to "sign" their review (i.e. to associate their name with their comments). Otherwise, all review comments remain anonymous.
  • All reviews of published articles are published. This includes manuscript files, peer review comments, author rebuttals and revised materials.
  • Each time a decision is made by the Academic Editor, each reviewer will receive a copy of the Decision Letter (which will include the comments of all reviewers).

If you have any questions about submitting your review, please email us at [email protected].