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Immunolocalization of GLDP in bundle sheath and mesophyll cells of (A, B) Homolepis aturensis, (C, D) Steinchisma laxum, and (E, F) S. hians. C, chloroplasts; p, peroxisomes; black asterisk, mitochondria; white asterisk, mitochondria surrounded by chloroplast. Scale bars=500 nm.  

Immunolocalization of GLDP in bundle sheath and mesophyll cells of (A, B) Homolepis aturensis, (C, D) Steinchisma laxum, and (E, F) S. hians. C, chloroplasts; p, peroxisomes; black asterisk, mitochondria; white asterisk, mitochondria surrounded by chloroplast. Scale bars=500 nm.  

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Photorespiratory glycine shuttling and decarboxylation in bundle sheath (BS) cells exhibited by C2 species is proposed to be the evolutionary bridge to C4 photosynthesis in eudicots. To evaluate this in grasses, we compare anatomy, cellular localization of glycine decarboxylase (GDC), and photosynthetic physiology of a suspected C2 grass, Homolepis...

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... observed for H. aturensis and S. hians, a significantly greater number of mitochondria and peroxisomes are positioned at the centrip- etal BS pole in S. laxum, although chloroplasts are arranged equally around BS cells (Fig 1C, D; Supplementary Table S4). BS mitochondria are commonly surrounded by chloro- plasts in S. hians (Fig 3E), H. aturensis, and S. laxum in a pattern similar to previous reports for Steinchisma species (Brown et al., 1983). In contrast to centripetal chloroplasts displayed with their long axis parallel to the BS wall in S. laxum, the long axis of these chloroplasts in H. aturensis and S. hians are perpendicular to the BS wall (Figs 1, 2). ...
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... peroxisomes, and chloroplasts are abundant in mestome sheath cells of N. minor (Hattersley et al., 1986;Sage et al., 2014). Unlike H. aturensis, S. hians, and S. laxum, organelles are not polarized in their distribu- tion ( Supplementary Fig. S3), but are instead positioned around the mestome sheath cell periphery (Hattersley et al., 1986;Sage et al., 2014). ...
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... from quantification of gold particles conjugated to secondary antibodies that bind to anti-GLDP are sum- marized in Supplementary Table S5. GLDP is almost exclusively located in BS cells of H. aturensis and S. hians, and mestome sheath cells of N. minor (Fig. 3A, B, E, F; Supplementary Fig. S3). Both BS and M cells contained high levels of GLDP labeling in S. laxum (Fig. 3C, D). In com- parison with all other C 3 species examined, S. laxum had the highest GLDP labeling in BS mitochondria (Supplementary Table S5). These patterns in GLDP distribution in M and BS mitochondria of S. laxum and S. ...
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... from quantification of gold particles conjugated to secondary antibodies that bind to anti-GLDP are sum- marized in Supplementary Table S5. GLDP is almost exclusively located in BS cells of H. aturensis and S. hians, and mestome sheath cells of N. minor (Fig. 3A, B, E, F; Supplementary Fig. S3). Both BS and M cells contained high levels of GLDP labeling in S. laxum (Fig. 3C, D). ...
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... of gold particles conjugated to secondary antibodies that bind to anti-GLDP are sum- marized in Supplementary Table S5. GLDP is almost exclusively located in BS cells of H. aturensis and S. hians, and mestome sheath cells of N. minor (Fig. 3A, B, E, F; Supplementary Fig. S3). Both BS and M cells contained high levels of GLDP labeling in S. laxum (Fig. 3C, D). In com- parison with all other C 3 species examined, S. laxum had the highest GLDP labeling in BS mitochondria (Supplementary Table S5). These patterns in GLDP distribution in M and BS mitochondria of S. laxum and S. hians are similar to earlier reports on these species (Hylton et al., 1988). The gold density in C 3 species D. ...
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... sheath cell ultrastructure of C 3 and C 4 species. Figure S3. Leaf structure and anatomy and immunolocali- zation of GLDP in N. minor. ...

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... However, one of the main bottlenecks is an incomplete understanding of how bundle sheath cells become photosynthetically activated in C 4 plants. On average, the bundle sheath chloroplast content of C 4 species is ~ 30% more than in C 3 species [16,18], but how this evolved is not fully understood. The GOLDEN2-LIKE family of transcription factors known to regulate chloroplast development in C 4 species [19][20][21]. ...
... However, the bundle sheath has been challenging to phenotype in C 3 plants. Classical bright-field light microscopy after embedding samples in resin and thin sectioning has been used [18]. Although this is simple and easily available, it only captures two-dimensional (2D) information from a thin section. ...
... It has been challenging to phenotype bundle sheath tissue in C 3 species as these cells are deeper in the leaf because of the many layers of mesophyll cells [27]. Approaches including bright-field light microscopy [18], transmission electron microscopy [23], serial block-face scanning electron microscopy [26] and single-cell isolation methods [22] are slow and so this hinders rapid analysis of transgenic lines harbouring candidate genes that are hypothesized to control chloroplast proliferation in the bundle sheath. ...
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... Shifting the glycine decarboxylation step exclusively to the bundle sheath cells leads to increased CO2 release in these cells, creating an elevated CO 2 environment around the bundle sheath Rubisco and considerably reducing the oxygenase reaction in this compartment. The bundle sheath specific localisation of the Pprotein from the glycine decarboxylase complex has been shown in C3-C4 species from diverse phylogenetic backgrounds by immunolocalization (Rawsthorne et al., 1988, Schlüter and Weber, 2016, Khoshravesh et al., 2016, Oono et al., 2022. The glycine shuttle biochemistry is accompanied by enhanced centripetal organelle accumulation in the bundle sheath cells (for reviews see: Weber, 2016, Lundgren, 2021). ...
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... Arthropogoninae (Khoshravesh et al., 2016) and Chenopodiaceae (Yorimitsu et al., 2019) 824 have shown a close arrangement of mitochondria and chloroplasts. Thus, BS ultrastructure 825 seems to play a major role in prevention of photorespiratory CO2 and NH3 loss and in 826 improvement of leaf carbon and possibly also nitrogen economy. ...
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