# Stromules: An Incident or Formalized Pathway for Molecules Transfer Between Organelles?

**Authors:** Joanna Grzyb

PMC · DOI: 10.3390/ijms262110680 · 2025-11-02

## TL;DR

This paper explores whether stromules, tubular extensions of chloroplasts, are involved in molecule transfer between organelles or are just incidental structures.

## Contribution

The paper provides a perspective on stromules' potential role in organelle communication and challenges their functional significance.

## Key findings

- Stromules are formed by the chloroplast envelope and are associated with the cytoskeleton.
- Stromules appear under illumination and after pathogen attack, suggesting a role in signaling.
- There is no consensus on whether stromules facilitate molecule transport or are functionally insignificant.

## Abstract

The stromules are tubular extensions of chloroplasts, or broader plastids, formed by the organelle envelope and filled with the stroma, the internal content of organelle. The formation of stromules is related to the cytoskeleton. Stromules occur in photosynthetic tissues under illumination and are therefore proposed to be important for retrograde signaling, which is essential for adaptation to stress factors. Stromules are also observed after pathogen attack. Some groups propose that stromules are a resemblance to endoplasmic reticulum dynamics, without having actual significance in molecules transport within a cell. However, there is no consensus among researchers regarding the actual function and significance of stromules, which can be the result of different models used to study stromules, and the necessity of using fluorescent labels, with all advantages and limitations of fluorescence-based methodology. Here I briefly discuss current knowledge on the subject from a perspective of stromule origin—the chloroplast envelope, and the potential advantages of having a conduit within organelles instead of relying on diffusion through cytosol.

## Full-text entities

- **Genes:** OEP37 (chloroplast outer envelope protein 37) [NCBI Gene 819000] {aka ARABIDOPSIS CHLOROPLAST OUTER ENVELOPE PROTEIN 37, ATOEP37, F6E13.8, chloroplast outer envelope protein 37}, OEP16-1 (outer plastid envelope protein 16-1) [NCBI Gene 817439] {aka ATOEP16-1, ATOEP16-L, F8N16.19, F8N16_19, OEP16, OUTER PLASTID ENVELOPE PROTEIN 16-L}, TLR4 (toll like receptor 4) [NCBI Gene 7099] {aka ARMD10, CD284, TLR-4, TOLL}, GUN1 (s uncoupled 1) [NCBI Gene 817698] {aka T28P16.11, T28P16_11, genomes uncoupled 1}, NRIP1 (nuclear receptor interacting protein 1) [NCBI Gene 8204] {aka CAKUT3, RIP140}, SAL1 (SAL1 phosphatase-like protein) [NCBI Gene 836519] {aka ALTERED EXPRESSION OF APX2 8, ALX8, ATSAL1, AtFRY1, FIERY1, FRY1}, BAK1 (BCL2 antagonist/killer 1) [NCBI Gene 578] {aka BAK, BAK-LIKE, BCL2L7, CDN1}, PADI4 (peptidyl arginine deiminase 4) [NCBI Gene 23569] {aka PAD, PAD4, PADI5, PDI4, PDI5}, ATI1 (mesoderm induction early response protein) [NCBI Gene 819206] {aka ATG8-interacting protein 1, F4I18.37, F4I18_37}, APE1 (acclimation of photosynthesis to environment) [NCBI Gene 833856] {aka ACCLIMATION OF PHOTOSYNTHESIS TO  ENVIRONMENT, ACCLIMATION OF PHOTOSYNTHESIS TO ENVIRONMENT, MBB18.21, MBB18_21}
- **Diseases:** infection (MESH:D007239), EDS1 (OMIM:607836), injury to (MESH:D014947), swelling (MESH:D004487)
- **Chemicals:** polyunsaturated fatty acids (MESH:D005231), MGDG (-), hydrogen peroxide (MESH:D006861), methyl jasmonate (MESH:C072239), fatty acid (MESH:D005227), starch (MESH:D013213), heme (MESH:D006418), lipid (MESH:D008055), Sucrose (MESH:D013395), PAP (MESH:D010724), mannitol (MESH:D008353), fructose (MESH:D005632), reactive oxygen species (MESH:D017382), glucose (MESH:D005947), DAPI (MESH:C007293), salicylic acid (MESH:D020156), ammonia (MESH:D000641), sugar (MESH:D000073893), phosphate (MESH:D010710), abscisic acid (MESH:D000040), spectinomycin (MESH:D000198), strigolactones (MESH:C000591191), streptomycin (MESH:D013307), chlorophyll (MESH:D002734), ethylene (MESH:C036216), isoprenoids (MESH:D013729), carotenoids (MESH:D002338), luminal (MESH:D010634), sorbitol (MESH:D013012), water (MESH:D014867), tetrapyrrole (MESH:D045725)
- **Species:** Agrobacterium (genus) [taxon 357], Arabidopsis thaliana (mouse-ear cress, species) [taxon 3702], Solanum lycopersicum (tomato, species) [taxon 4081], Homo sapiens (human, species) [taxon 9606], Nicotiana benthamiana (species) [taxon 4100], Nicotiana tabacum (American tobacco, species) [taxon 4097]

## Figures

3 figures with captions in the complete paper: https://tomesphere.com/paper/PMC12608775/full.md

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Source: https://tomesphere.com/paper/PMC12608775