1. Wada, H., Hirasawa, R., Omata, T. and Murata, N.

(1984) The lipid phase of

thylakoid and cytoplasmic membranes

from the blue-green algae

(cyanobacteria), Anacystis nidulans and Anabaena variabilis. Plant Cell Physiol.

25:907-911.
2. Murata, N., Wada, H. and Hirasawa, R. (1984) Reversible and irreversible
inactivation of photosynthesis in relation to the lipid phase of membranes in the
blue-green algae (cyanobacteria), Anacystis nidulans and Anabaena variabilis.
Plant Cell Physiol. 25:1027-1032.
3. Gonda, K., Wada, H. and Murase, N. (1985) Calorimetric studies of glycolysis of E.
coli under non-growing condition. Thermochim. Acta 88:339-344.
4. Wada, H. and Murata, N. (1989) Synechocystis PCC6803 mutants defective in
desaturation of fatty acids. Plant Cell Physiol. 30:971-978.
5. Wada, H. and Murata, N. (1990) Temperature-induced changes in fatty acid
composition in the cyanobacterium, Synechocystis PCC6803. Plant Physiol.
92:1062-1069.
6. Wada, H., Gombos Z. and Murata, N. (1990) Enhancement of chilling tolerance of
a cyanobacterium by genetic manipulation of fatty acid desaturation. Nature
347:200-203.
7. Gombos, Z., Wada, H. and Murata, N. (1991) Direct evaluation of effects of
fatty-acid unsaturation on the thermal properties of photosynthetic activity, as
studied by mutation and transformation of Synechocystis PCC6803. Plant Cell
Physiol. 32:205-211.
8. Mamedov, M. D., Hayashi, H., Wada, H., Mohanty, P. S., Papageorgiou, G. C. and
Murata, N. (1991) Glycinebetaine enhances and stabilizes the evolution of oxygen
and the synthesis of ATP by cyanobacterial thylakoid membranes, FEBS Lett.
294:271-274.

9. Lehel, C., Wada, H., Kovacs, E., Torok, Z., Gombos, Z., Horvath, I., Murata, N.
and Vigh, L. (1992) Heat shock protein synthesis of the cyanobacterium
Synechocystis PCC6803: Purification of the groEL-related chaperonin. Plant Mol.
Biol. 18:327-336.
10. Wada, H., Gombos, Z., Sakamoto, T. and Murata, N. (1992) Genetic manipulation
of the extent of desaturation of fatty acids in membrane lipids in the
cyanobacterium Synechocystis PCC6803. Plant Cell Physiol. 33:535-540.
11. Murata, N., Wada, H. and Gombos, Z. (1992) Modes of fatty-acid desaturation in
cyanobacteria. Plant Cell Physiol. 33:933-941.
12. Gombos, Z., Wada, H. and Murata, N. (1992) Unsaturation of fatty acids in
membrane lipids enhances the tolerance of the cyanobacterium Synechocystis
PCC6803 to low-temperature photoinhibition. Proc. Natl. Acad. Sci. USA
89:9959-9963.
13. Wada, H., Schmidt, H., Heinz, E. and Murata, N. (1993) In vitro
ferredoxin-dependent desaturation of fatty acids in cyanobacterial thylakoid
membranes. J. Bacteriol. 175:544-547.
14. Lehel, C., Los, D. A., Wada, H., Grorgyey, A., Horvath, I., Kovacs, E., Murata, N.
and Vigh, L. (1993) A second groEL-like gene, organized in a groESL operon is
present in the genome of Synechocystis sp. PCC6803. J. Biol. Chem.
268:1799-1804.
15. Malakhov, M. P., Wada, H., Los, D. A., Sakamoto, T. and Murata, N. (1993)
Structure of the cyanobacterial gene for the 50S ribosomal protein L9. Plant Mol.
Biol. 21:913-918.

16. Wada, H., Avelange-Macherel, M.-H. and Murata, N. (1993) The desA gene of the
cyanobacterium Synechocystis PCC6803 is the structural gene for 12 desaturase. J.
Bacteriol. 175:5056-5058.
17. Gombos, Z., Wada, H., Hideg, E. and Murata, N. (1994) The unsaturation of
membrane lipids stabilizes photosynthesis against heat stress. Plant Physiol.
104:563-567.
18. Sakamoto, T., Wada, H., Nishida, I., Ohmori, M. and Murata, N. (1994)
Identification of conserved domains in the 12 desaturases of cyanobacteria. Plant
Mol. Biol. 24:643-650.
19. Wada, H., Gombos, Z. and Murata, N. (1994) Contribution of membrane lipids to
the ability of the photosynthetic machinery to tolerate temperature stresses. Proc.
Natl. Acad. Sci. USA 91:4273-4277.
20. Malakhov, M. P., Wada, H., Los, D. A. and Murata, N. (1994) A new type of
cytochrome c from Synechocystis PCC6803. J. Plant Physiol. 144:259-264.
21. Sakamoto, T., Higashi, S., Los, D., Wada, H., Nishida, I., Ohmori, M. and Murata,
N. (1994) Cloning of 3 desaturase from cyanobacteria and its use in altering the
degree of membrane lipid unsaturation. Plant Mol. Biol. 26:249-263.
22. Gombos, Z., Wada, H. and Murata, N. (1994) The recovery of photosynthesis from
low-temperature photoinhibition is accelerated by the unsaturation of membrane
lipids: A mechanism of chilling tolerance. Proc. Natl. Acad. Sci. USA 91:8787-8791.
23. Sakamoto, T., Wada, H., Nishida, I., Ohmori, M. and Murata, N. (1994) -9
acyl-lipid desaturases of cyanobacteria: Molecular cloning and substrate
specificities in terms of fatty acids, sn-positions, and polar head group. J. Biol.
Chem. 41:25576-25580.

24. Malakhov, M. P., Los, D. A., Wada, H., Semenenko, V. E. and Murata, N. (1995)
Characterization of the murF gene of the cyanobacterium Synechocystis PCC6803.
Microbiology 141:163-169.
25. Avelange-Macherel, M.-H., Macherel, D., Wada, H. and Murata, N. (1995)
Site-directed mutagenesis of histidine residues in the 12 acyl-lipid desaturase of
Synechocystis. FEBS Lett. 361:111-114.
26. Gombos, Z., Hideg, E., Asiros, O., Wada, H. and Murata, N. (1995) The role of
lipid desaturation in protection mechanism against temperature stresses. Acta
Phytopathol. Entmol. Hung. 30:89-92.
27. Murata, N. and Wada, H. (1995) Acyl-lipid desaturases and their importance in the
tolerance and acclimation to cold of cyanobacteria. Biochem. J. 308:1-8.
28. Gombos, Z., Wada, H., Varkonyi, Z., Sakamoto, T. and Murata, N. (1996)
Characterization of the Fad12 mutant of Synechocystis that is defective in 12
acyl-lipid desaturase activity. Biochim. Biophys. Acta 1299:117-123.
29. Malakhov, M., Wada, H., Los, D. and Murata, N. (1996) The coxD gene for heme O
synthase in Synechocystis. Biochim. Biophys. Acta 1273:84-86.
30. Wada, H., Shintani, D. and Ohlrogge, J. (1997) Why do mitochondria synthesize
fatty acids? Evidence for involvement in lipoic acid production. Proc. Natl. Acad.
Sci. USA 94:1591-1596.
31. Sakamoto, T., Higashi, S., Wada, H., Murata, N. and Bryant, D. A. (1997)
Low-temperature induced desaturation of fatty acids and expression of desaturase
genes in the cyanobacterium Synechococcus sp. PCC 7002. FEMS Microbiol. Lett.
152:313-320.

32. Okumura, N., Masamoto, K. and Wada, H. (1997) The gshB gene in the
cyanobacterium Synechococcus sp. PCC 7942 encodes a functional glutathione
synthetase. Microbiology 143:2883-2890.
33. Aoki, S., Kondo, T., Wada, H. and Ishiura M. (1997) Circadian rhythm of the
cyanobacterium Synechocystis sp. PCC 6803 in the dark. J. Bacteriol.
179:5751-5755.
34. Kis, M., Zsiros, O., Farkas, T., Wada, H., Nagy, F. and Gombos, Z. (1998)
Light-induced expression of fatty acid desaturase genes. Proc. Natl. Acad. Sci. USA
95:4209-4214.
35. Yasuno, R. and Wada, H. (1998) Biosynthesis of lipoic acid in Arabidopsis thaliana.
Cloning and characterization of the cDNA for lipoic acid synthase. Plant Physiol.
118:935-943.
36. Hagio, M., Gombos, Z., Vrkonyi, Z., Masamoto, K., Sato, N., Tsuzuki, M. and
Wada, H. (2000) Direct evidence for requirement of phosphatidylglycerol in
photosystem II of photosynthesis. Plant Physiol. 124:795-804.
37. Sato, N., Hagio, M., Wada, H. and Tsuzuki, M. (2000) Environmental effects on
acidic lipids of thylakoid membranes. Biochem. Soc. Trans. 28:912-914.
38. Sato, N., Hagio, M., Wada, H. and Tsuzuki, M. (2000) Requirement of
phosphatidylglycerol for photosynthetic function in thylakoid membranes. Proc.
Natl. Acad. Sci. USA 97:10655-10660.
39. Wada, M., Yasuno, R., Jordan, S. W., Cronan, J. E., Jr., Wada, H. (2001) Lipoic
acid metabolism in Arabidopsis thaliana: Cloning and characterization of a cDNA
encoding lipoyltransferase. Plant Cell Physiol. 42:650-656.

40. Morikawa, T., Yasuno, R. and Wada, H. (2001) Do mammalian cells synthesize
lipoic acid? Identification of a mouse cDNA for lipoic acid synthase located in
mitochondria. FEBS Lett. 498:16-21.
41. Wada, M., Yasuno, R. and Wada, H. (2001) Identification of an Arabidopsis cDNA
for lipoyltransferase located in plastids. FEBS Lett. 506:286-290.
42. Masamoto, K., Wada, H., Kaneko, T. and Takaichi, S. (2001) Identification of a
gene required for cis-to-trans carotene isomerization in carotenogenesis of the
cyanobacterium Synechocystis sp. PCC6803. Plant Cell Physiol. 42:1398-1402.
43. Gombos, Z., Vrkonyi, Z., Hagio, M., Iwaki, M., Kovcs, L., Masamoto, K., Itoh, S.
and Wada, H. (2002) Phosphatidylglycerol requirement for the function of electron
acceptor plastoquinone QB in the photosystem II reaction center. Biochemistry
41:3796-3802.
44. Vrkonyi, Z., Masamoto, K., Debreczeny, M., Zsiros, O., Ughy, B., Gombos, Z.,
Domonkos, I., Farkas, T., Wada, H. and Szalontai, B. (2002)
Low-temperature-induced accumulation of xanthophylls and its structural
consequences in the photosynthetic membranes of the cyanobacterium
Cylindrospermopsis raciborskii: An FT-IR spectroscopic study. Proc. Natl. Acad.
Sci. USA 99:2410-2415.
45. Xu, C., Hrtel, H., Wada, H., Hagio, M., Yu, B., Eakin, C. and Benning, C. (2002)
The pgp1 mutant locus of Arabidopsis encodes a phosphatidylglycerolphosphate
synthase with impaired activity. Plant Physiol. 129:594-604.
46. Yasuno, R. and Wada, H. (2002) The biosynthetic pathway for lipoic acid is present
in plastids and mitochondria in Arabidopsis thaliana. FEBS Lett. 517:110-114.
47. Hagio, M., Sakurai, I., Sato, S., Kato, T., Tabata, S. and Wada, H. (2002)
Phosphatidylglycerol is essential for the development of thylakoid membranes in
Arabidopsis thaliana. Plant Cell Physiol. 43:1456-1464.

48. Sakurai, I., Hagio, M., Gombos, Z., Tyystjrvi, T., Paakkarinen, V., Aro, E.-M. and
Wada, H. (2003) Requirement of phosphatidylglycerol for maintenance of
photosynthetic machinery. Plant Physiol. 133:1376-1384.
49. Yasuno, R., von Wettstein-Knowles, P. and Wada, H. (2004) Identification and
molecular characterization of -ketoacyl-[acyl carrier protein] synthase of
Arabidopsis thaliana involved in mitochondrial fatty acid synthesis. J. Biol. Chem.
279:8242-8251.
50. Domonkos, I, Malec, P., Sallai, A., Kovcs, L., Itoh, K., Shen, G., Ughy, B., Bogos,
B., Sakurai, I., Kis, M., Strzalka, K., Wada, H., Itoh, S., Farkas, T. and Gombos, Z.
(2004) Phosphatidylglycerol is essential for oligomerization of PS I reaction center.
Plant Physiol. 134:1471-1478.
51. Masamoto, K., Hisatomi, S., Sakurai, I., Gombos, Z. and Wada, H. (2004)
Requirement of carotene isomerization for the assembly of photosystem II in
Synechocystis sp. PCC6803. Plant Cell Physiol. 45:1325-1329.
52. Katayama, K., Sakurai, I. and Wada, H. (2004) Identification of an Arabidopsis
thaliana gene for cardiolipin synthase located in mitochondria. FEBS Lett.
577:193-198.
53. Awai, K., Kakimoto, T., Awai, C., Kaneko, T., Nakamura, Y., Takamiya, K., Wada,
H. and Ohta H. (2006) Comparative genomic analysis revealed a gene for
monoglucosyldiacylglycerol synthase, an enzyme for photosynthetic membrane
lipid synthesis in cyanobacteria. Plant Physiol. 141:1120-1127.
54. Sakurai, I., Shen, J.-R. Jing, Leng, Ohashi, S., Kobayashi, M. and Wada, H. (2006)
Lipids in oxygen-evolving photosystem II complexes of cyanobacteria and higher
plants. J. Biochem. 140:201-209.

55. Wada, H. and Murata, N. (2007) The essential role of phosphatidylglycerol in

photosynthesis. Photosynth. Res. 92:205-215.
56. Sakurai, I., Mizusawa, N., Ohashi, S., Kobayashi, M. and Wada, H. (2007) Effects
of the lack of phosphatidylglycerol on the donor side of photosystem II. Plant
Physiol. 144:1336-1346.
57. Sakurai, I., Mizusawa, N., Wada, H. and Sato, N. (2007) Digalactosyldiacylglycerol
is required for stabilization of oxygen-evolving complex in photosystem II. Plant
Physiol. 145:1361-1370.
58. Otsubo, M., Inoue, S., Tasaka, Y., Saito, A., Yasuno, R., Ikoma, C., Nagahata, Y.,
Tamura, N. and Wada, H. (2007) Isolation of freezing-tolerant mutants of
Arabidopsis thaliana by activation tagging. Cryobiol. Cryotech. 53:75-85.
59. Ujihara, T., Sakurai, I., Mizusawa, N. and Wada, H. (2008) A method for analyzing
lipid-modified proteins with mass spectrometry. Anal. Biochem. 374:429-431.
60. Apostolova, E. L., Domonkos, I., Dobrikova, A. G., Sallai, A., Bogos, B., Wada, H.,
Gombos, Z. and Taneva, S. G. (2008) Effect of phosphatidylglycerol depletion on
the surface electric properties and energy distribution in thylakoid membranes. J.
Photochem. Photobiol. 91:41-50.
61. Leng, J., Sakurai, I., Wada, H. and Shen J.-R. (2008) Effects of phospholipases and
lipase treatments on photosystem II core dimer from a thermophilic cyanobacterium.
Photosynth. Res. 98:469-478.
62. Mizusawa, N., Sakurai, I., Sato, N. and Wada, H. (2009) Lack of
digalactosyldiacylglycerol increases the sensitivity of Synechocystis sp. PCC 6803 to
high light stress. FEBS Lett. 583:718-722.
63. Domonkos, I., Malec, P., Laczko-Dobos, H., Bozkurt, O., Klodawska, K., Wada, H.,
Strzalka, K. and Gombos, Z. (2009) Phosphatidylglycerol depletion induces an

increase in myxoxanthophyll biosynthetic activity in Synechocystis PCC 6803 cells.

Plant Cell Physiol. 50:374-382.
64. Katayama, K. and Wada, H. (2009) Cellular function of the phospholipid cardiolipin.
Plant Morphol. 21:17-28.
65. Mizusawa, N., Sakata, S., Sakurai, I., Sato, N. and Wada, H. (2009) Involvement of
digalactosyldiacylglycerol in cellular thermotolerance of Synechocystis sp. PCC
6803. Arch. Microbiol. 191:595-601.
66. Bogos, B., Ughy, B., Domonkos, I., Laczk-Dobos, H., Komenda, J., Abasova, L.,
Cser, K., Vass, I., Sallai, A., Wada, H. and Gombos, Z. (2010) Phosphatidylglycerol
depletion affects photosystem II activity in Synechococcus PCC 7942 cells.
Photosynth. Res. 103:19-30.
67. Kubota, H., Sakurai, I., Katayama, K., Mizusawa, N., Ohashi, S., Kobayashi, M.,
Zhang, P., Aro, E.-M. and Wada, H. (2010) Purification and characterization of PSI
complex from Synechocystis sp. PCC 6803 by expressing histidine-tagged subunits.
Biochim. Biophys. Acta 1797:98-105.
68. Laczko-Dobos, H., Fryck, P., Ughy, B., Domonkos, I., Wada, H., Prokai, L. and
Gombos, Z. (2010) Remodeling of phosphatidylglycerol in Synechocystis PCC6803.
Biochim. Biophys. Acta 1801:163-170.
69. Nanjo, Y., Mizusawa, N., Wada, H., Slabas, A. R., Hayashi, H. and Nishiyama, Y.
(2010) Synthesis of fatty acids de novo is required for the photosynthetic acclimation
of Synechocystis sp. PCC 6803 to high temperature. Biochim. Biophys. Acta
1797:1483-1490.
70. Watanabe, M., Kubota, H., Wada, H., Narikawa, R. and Ikeuchi, M. (2011) Novel
supercomplex organization of Photosystem I in Anabaena and Cyanophora paradoxa. Plant

Cell Physiol. 52:162-168.

71. Mizusawa, N. and Wada, H. (2012) The role of lipids in photosystem II. Biochim.
Biophys. Acta 1817:194-208.
72. Katayama, K and Wada, H. (2012) T-DNA insertion in the CLS gene for cardiolipin
synthase affects development of Arabidopsis thaliana. Cytologia 77:123-129.
73. Chan, T., Shimizu, Y., Pospil, P., Nijo, N., Fujiwara, A., Taninaka, Y., Hori, H.,
Ishikawa, T., Nanba, D., Imai, A., Morita, N., Yoshioka-Nishimura, M., Izumi, Y.,
Yamamoto, Y., Kobayashi, H., Mizusawa, N., Wada, H. and Yamamoto, Y. (2012)
Quality control of photosystem II. Lipid peroxidation causes subsidiary but
substantial damage to photosystem II under illumination. PLoS ONE 7:e52100.
74. Kobayashi, K., Narise, T., Sonoike, K., Hashimoto, H., Sato, N., Kondo, M.,
Nishimura, M., Sato, M., Toyooka, K., Sugimoto, K., Wada, H., Masuda, T. and
Ohta, H. (2013) Role of galactolipid biosynthesis in coordinated development of
photosynthetic complexes and thylakoid membranes during chloroplast biogenesis in
Arabidopsis. Plant J. 73:250-261.
75. Sakata, S., Mizusawa, N., Kubota-Kawai, H., Sakurai, I. and Wada H. (2013) Psb28
is involved in recovery of photosystem II at high temperature in Synechocystis sp.
PCC 6803. Biochim. Biophys. Acta 1827:50-59.
76. Kobayashi, K., Sasaki, D., Noguchi, K., Fujinuma, D., Komatsu, H., Kobayashi, M.,
Sato, M., Toyooka, K., Sugimoto, K., Niyogi, K. K., Wada, H. and Masuda, T.
(2013) Photosynthesis in root chloroplasts developed in Arabidopsis lines
overexpressing GOLDEN2-LIKE transcription factors. Plant Cell Physiol.
54:1365-1377.
77. Hakkila, K., Antal, T., Rehman, A. U., Kurkela, J., Wada, H., Vass, I., Tyystjrvi, E.
and Tyystjrvi, T. (2014) Oxidative stress and photoinhibition can be separated in
the cyanobacterium Synechocystis sp. PCC 6803. Biochim. Biophys. Acta
1837:217-225.

78. Yuzawa Y., Shimojima, M., Sato, R., Mizusawa, N., Yamamichi, K., Suzuki, M.,
Iwai, M., Hori, K., Wada, H., Masuda, S. and Ohta, H. (2014) Cyanobacterial
monogalactosyldiacylglycerol-synthesis pathway is involved in normal unsaturation
of galactolipids and low-temperature adaptation of Synechocystis sp. PCC 6803.
Biochim. Biophys. Acta 1841:475-483.
79. Gunnelius, L., Hakkila, K., Kurkela, J., Wada, H., Tyystjrvi, E. and Tyystjrvi, T.
(2014) The omega subunit of the RNA polymerase core directs transcription
efficiency in cyanobacteria. Nucleic Acids Res. 42:4606-4614.
80. Tanoue, R., Kobayashi, M., Katayama, K., Nagata, N. and Wada, H. (2014)
Phosphatidylglycerol biosynthesis is required for the development of embryos and
normal membrane structures of chloroplasts and mitochondria in Arabidopsis. FEBS
Lett. 588:1680-1685.
81. Hori, K., Maruyama, F., Fujisawa, T., Togashi, T., Yamamoto, N., Seo, M., Sato, S.,
Yamada, T., Mori, H., Tajima, N., Moriyama, T., Ikeuchi, M., Watanabe, M., Wada,
H., Kobayashi, K., Saito, M., Masuda, T., Sasaki-Sekimoto, Y., Mashiguchi, K.,
Awai, K., Shimojima, M., Masuda, S., Iwai, M., Nobusawa, T., Narise, T., Kondo, S.,
Saito, H., Sato, R., Murakawa, M., Ihara, Y., Oshima, Y., Ohtaka, K., Satoh, M.,
Sonobe, K., Ishii, M., Ohtani, R., Kanamori, M., Honoki, R., Miyazaki, D.,
Mochizuki, H., Umetsu, J., Higashi, K., Shibata, D., Kamiya, Y., Sato, N., Nakamura,
Y., Tabata, S., Ida, S., Kurokawa, K. and Ohta, H. (2014) Klebsormidium flaccidum
genome reveals primary factors for plant terrestrial adaptation. Nat. Commun.
5:3978.
82. Kobayashi, K., Fujii, S., Sasaki, D., Baba, S., Ohta, H., Masuda, T. and Wada, H.
(2014) Transcriptional regulation of thylakoid galactolipid biosynthesis with
chlorophyll biosynthesis during chloroplast development in Arabidopsis. Front.
Plant Sci. 5:272.
83. Ujihara, T., Nagano, M., Wada, H. and Mitsuhashi, S. (2014) Identification of a
novel type of polyunsaturated fatty acid synthase involved in the biosynthesis of

arachidonic acid. FEBS Lett. 588:4032-4036.

84. Kobayashi, K., Masuda, T., Tajima, N., Wada, H. and Sato, N. (2014) Molecular
phylogeny and intricate evolutionary history of the three isofunctional enzymes
involved in the oxidation of protoporphyrinogen IX. Genome Biology and Evolution.
6:2141-2155
85. Fujii, S., Kobayashi, K., Niyogi, K. K., Nakamura, N. and Wada, H. (2014)
Inducible knockdown of the MGD1 gene for monogalactosyldiacylglycerol synthase
1 reveals roles of galactolipid biosynthesis during initial growth in Arabidopsis
seedlings. Plant Physiol. 166:1436-1449.
86. Kobayashi, K., Fujii, S., Sato, M., Toyooka, K. and Wada, H. (2015) Specific role
of phosphatidylglycerol and functional overlaps with other thylakoid lipids in
Arabidopsis chloroplast biogenesis. Plant Cell Rep. 34:631-642.
87. Okazaki, K., Miyagishima, S. and Wada, H. (2015) Phosphatidylinositol
4-phosphate negatively regulates chloroplast division. Plant Cell 27:663-674.
88. Hung, C.-H., Kobayashi, K., Wada, H. and Nakamura, Y. (2015) Isolation and
characterization of a phosphatidylglycerophosphate phosphatase1, PGPP1, in
Chlamydomonas reinhardtii. Plant Physiol. Biochem. 92:56-61.
89. Endo, K., Mizusawa, N., Shen, J.-R., Yamada, M., Tomo, T., Komatsu, H.,
Kobayashi, M., Kobayashi, K. and Wada, H. (2015) Site-directed mutagenesis of
amino acid residues of D1 protein interacting with phosphatidylglycerol affects the
function of plastoquinone QB in photosystem II. Accepted to Photosynth. Res.