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The structure of the Physcomitrium patens photosystem I reveals a unique Lhca2 paralogue replacing Lhca4

Nature Plants volume 8, pages 307–316 (2022)Cite this article

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Abstract

The moss Physcomitrium patens diverged from green algae shortly after the colonization of land by ancient plants. This colonization posed new environmental challenges, which drove evolutionary processes. The photosynthetic machinery of modern flowering plants is adapted to the high light conditions on land. Red-shifted Lhca4 antennae are present in the photosystem I light-harvesting complex of many green-lineage plants but absent in P. patens. The cryo-EM structure of the P. patens photosystem I light-harvesting complex I supercomplex (PSI–LHCI) at 2.8 Å reveals that Lhca4 is replaced by a unique Lhca2 paralogue in moss. This PSI–LHCI supercomplex also retains the PsaM subunit, present in Cyanobacteria and several algal species but lost in vascular plants, and the PsaO subunit responsible for binding light-harvesting complex II. The blue-shifted Lhca2 paralogue and chlorophyll b enrichment relative to flowering plants make the P. patens PSI–LHCI spectroscopically unique among other green-lineage supercomplexes. Overall, the structure represents an evolutionary intermediate PSI with the crescent-shaped LHCI common in vascular plants, and contains a unique Lhca2 paralogue that facilitates the moss’s adaptation to low-light niches.

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Fig. 1: The overall structure of the moss PSI–LHCI complex.
Fig. 2: The P. patens Lhca2b occupies the plant Lhca4 position in the LHCI antenna.
Fig. 3: Antennae specificity in LHCI.
Fig. 4: Structural differences in Chl coordination between higher plants and moss.
Fig. 5: Comparison of Chl b positions between P. patens and P. sativum PSI–LHCI.
Fig. 6: Red Chl content in the LHCI antenna in the green lineage.

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Data availability

The models and maps of the final model (PDB ID: 7SKQ; EMDB: EMD-23023); the PSI–LHCI model, excluding PsaO (PDB ID: 7KUX; EMDB: EMD-23040); and the PsaO alone (PDB ID: 7KU5; EMDB: EMD-23034) are all deposited in the Protein Databank and Electron Microscopy Database, respectively. Source data are provided with this paper.

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Acknowledgements

We thank Prof. N. Ohad and Dr R. Yaari for help with moss strains, cultivation and protocols; the Arizona State University for use of the Titan Krios at the Erying Materials Center; and the National Science Foundation (No. MRI 1531991) for funding of this instrument. This study was funded by a startup grant from Arizona State University and supported by grant number 2034021 from the National Science Foundation to Y.M.

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  1. These authors contributed equally: C. Gorski, R. Riddle, H. Toporik.

Authors and Affiliations

  1. School of Molecular Sciences, Arizona State University, Tempe, AZ, USA

    C. Gorski, R. Riddle, H. Toporik, Z. Da, Z. Dobson & Y. Mazor

  2. Biodesign Center for Applied Structural Discovery, Arizona State University, Tempe, AZ, USA

    C. Gorski, R. Riddle, H. Toporik, Z. Da, Z. Dobson & Y. Mazor

  3. John M. Cowley Center for High Resolution Electron Microscopy, Arizona State University, Tempe, AZ, USA

    D. Williams

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Contributions

C.G. and R.R. performed sample preparation and measurements. Z. Da and Z. Dobson performed measurements. D.W. performed cryo-EM data acquisition. Y.M., C.G. and R.R. analysed the data and performed structural modelling. C.G., H.T., R.R. and Y.M. planned the experiments and wrote the manuscript.

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Correspondence to Y. Mazor.

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Supplementary Figs. 1–7, uncropped blots for Fig. 5c and Tables 1–3.

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Source data

Source Data Fig. 1

Fig. 1d,e source data – absorption and emission data.

Source Data Fig. 5

Fig. 5c,d source data – HPLC chromatograms and absorption data.

Source Data Table 1

Table 1 source data – HPLC chromatograms.

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Gorski, C., Riddle, R., Toporik, H. et al. The structure of the Physcomitrium patens photosystem I reveals a unique Lhca2 paralogue replacing Lhca4. Nat. Plants 8, 307–316 (2022). https://doi.org/10.1038/s41477-022-01099-w

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