DOI: 10.1042/bj20101538. The light-harvesting complexes of higher-plant Photosystem I: Lhca1/4 and Lhca2/3 form two red-emitting heterodimers

The light-harvesting complexes of higher-plant Photosystem I: Lhca1/4 and Lhca2/3 form two red-emitting heterodimers

10.1042/bj20101538

Crossref journal-article

Portland Press Ltd.

Biochemical Journal (288)

Abstract

The outer antenna of higher-plant PSI (Photosystem I) is composed of four complexes [Lhc (light-harvesting complex) a1–Lhca4] belonging to the light-harvesting protein family. Difficulties in their purification have so far prevented the determination of their properties and most of the knowledge about Lhcas has been obtained from the study of the in vitro reconstituted antennas. In the present study we were able to purify the native complexes, showing that Lhca2/3 and Lhca1/4 form two functional heterodimers. Both dimers show red-fluorescence emission with maxima around 730 nm, as in the intact PSI complex. This indicates that the dimers are in their native state and that LHCI-680, which was previously assumed to be part of the PSI antenna, does not represent the native state of the system. The data show that the light-harvesting properties of the two dimers are functionally identical, concerning absorption, long-wavelength emission and fluorescence quantum yield, whereas they differ in their high-light response. Implications of the present study for the understanding of the energy transfer process in PSI are discussed. Finally, the comparison of the properties of the native dimers with those of the reconstituted complexes demonstrates that all of the major properties of the Lhcas are reproduced in the in vitro systems.

Bibliography

Wientjes, E., & Croce, R. (2011). The light-harvesting complexes of higher-plant Photosystem I: Lhca1/4 and Lhca2/3 form two red-emitting heterodimers. Biochemical Journal, 433(3), 477â485.

Authors 2

Emilie Wientjes (first)
Roberta Croce (additional)

References 56 Referenced 105

10.1038/nature05687 / Nature / The structure of a plant Photosystem I supercomplex at 3.4 Å resolution by Amunts (2007)
10.1038/35082000 / Nature / Three-dimensional structure of cyanobacterial Photosystem I at 2.5 Å resolution by Jordan (2001)
10.1007/978-94-011-3953-3_100 / Photosynthesis: Mechanisms and Effects / The light-harvesting complex of photosytem I: pigment composition and stoichiometry by Croce (1998)
10.1016/S1360-1385(99)01419-3 / Trends Plant Sci. / A guide to the Lhc genes and their relatives in Arabidopsis by Jansson (1999)
10.1021/bi0015358 / Biochemistry / Green plant Photosystem I binds light-harvesting complex I on one side of the complex by Boekema (2001)
10.1038/nature02200 / Nature / Crystal structure of plant Photosystem I by Ben-Shem (2003)
10.1111/j.1432-1033.2004.04426.x / Eur. J. Biochem. / Stoichiometry of LHCI antenna polypeptides and characterization of gap and linker pigments in higher plants Photosystem I by Ballottari (2004)
10.1074/jbc.M606417200 / J. Biol. Chem. / Contrasting behavior of higher plant Photosystem I and II antenna systems during acclimation by Ballottari (2007)
10.1186/1471-2229-9-71 / BMC Plant Biol. / Antenna complexes protect Photosystem I from photoinhibition by Alboresi (2009)
10.1111/j.1751-1097.2007.00136.x / Photochem. Photobiol. / Selective photobleaching of chlorophylls and carotenoids in Photosystem I particles under high-light treatment by Andreeva (2007)
10.1562/0031-8655(2000)072<0508:DOTPIC>2.0.CO;2 / Photochem. Photobiol. / Degradation of the Photosystem I complex during photoinhibition by Hui (2000)
10.1016/S0005-2728(01)00203-1 / Biochim. Biophys. Acta / Energy transfer and trapping in Photosystem I by Gobets (2001)
10.1023/A:1006236829711 / Photosyn. Res. / The importance of PSI chlorophyll red forms in light-harvesting by leaves by Rivadossi (1999)
10.1007/BF00016556 / Photosyn. Res. / Long-wavelength absorbing antenna pigments and heterogeneous absorption-bands concentrate excitons and increase absorption cross-section by Trissl (1993)
10.1016/j.bbabio.2004.11.007 / Biochim. Biophys. Acta / Kinetics of excitation trapping in intact Photosystem I of Chlamydomonas reinhardtii and Arabidopsis thaliana by Ihalainen (2005)
10.1016/S0005-2728(02)00399-7 / Biochim. Biophys. Acta / The photochemical trapping rate from red spectral states in PSI-LHCI is determined by thermal activation of energy transfer to bulk chlorophylls by Jennings (2003)
10.1021/bi960214m / Biochemistry / Excited state equilibration in the Photosystem I light-harvesting I complex: P700 is almost isoenergetic with its antenna by Croce (1996)
10.1529/biophysj.107.117101 / Biophys. J. / Trap-limited charge separation kinetics in higher plant Photosystem I complexes by Slavov (2008)
10.1529/biophysj.108.140467 / Biophys. J. / Picosecond fluorescence of intact and dissolved PSI-LHCI crystals by van Oort (2008)
10.1111/j.1432-1033.1987.tb10791.x / Eur. J. Biochem. / Chlorophyll–protein complexes of barley Photosystem-I by Bassi (1987)
10.1111/j.1432-1033.1992.tb16918.x / Eur. J. Biochem. / Identification of the Photosystem I antenna polypeptides in barley: isolation of three pigment-binding antenna complexes by Knoetzel (1992)
10.1016/0014-5793(84)80197-0 / FEBS Lett. / Chlorophyll a/b proteins of Photosystem I by Lam (1984)
10.1074/jbc.M205889200 / J. Biol. Chem. / Pigment binding of Photosystem I light-harvesting proteins by Schmid (2002)
10.1007/BF00032234 / Photosyn. Res. / Rapid isolation of Photosystem I chlorophyll-binding proteins by anion-exchange perfusion chromatography by Tjus (1995)
10.1021/bi9813227 / Biochemistry / A thermal broadening study of the antenna chlorophylls in PSI-200, LHCI, and PSI core by Croce (1998)
10.1021/bi0007369 / Biochemistry / Evidence for two spectroscopically different dimers of light-harvesting complex I from green plants by Ihalainen (2000)
{'key': '2021112214452763100_B27', 'first-page': '29', 'article-title': 'The Lhca antenna complexes of higher plants photosystem I. Biochim', 'volume': '1556', 'author': 'Croce', 'year': '2002', 'journal-title': 'Biophy. Acta'} / Biophy. Acta / The Lhca antenna complexes of higher plants photosystem I. Biochim by Croce (2002)
10.1021/bi027398r / Biochemistry / Recombinant Lhca2 and Lhca3 subunits of the Photosystem I antenna system by Castelletti (2003)
10.1073/pnas.94.14.7667 / Proc. Natl. Acad. Sci. U.S.A. / In vitro reconstitution of the Photosystem I light-harvesting complex LHCI-730: heterodimerization is required for antenna pigment organization by Schmid (1997)
10.1038/nature02373 / Nature / Crystal structure of spinach major light-harvesting complex at 2.72 Å resolution by Liu (2004)
10.1074/jbc.M408908200 / J. Biol. Chem. / Origin of the 701 nm fluorescence emission of the Lhca2 subunit of higher plant Photosystem I by Croce (2004)
10.1074/jbc.M309203200 / J. Biol. Chem. / The nature of a chlorophyll ligand in Lhca proteins determines the far red fluorescence emission typical of Photosystem I by Morosinotto (2003)
10.1074/jbc.M205062200 / J. Biol. Chem. / Mutation analysis of Lhca1 antenna complex: low energy absorption forms originate from pigment–pigment interactions by Morosinotto (2002)
10.1074/jbc.M500705200 / J. Biol. Chem. / Pigment-pigment interactions in Lhca4 antenna complex of higher plants Photosystem I by Morosinotto (2005)
10.1016/j.bbabio.2006.06.018 / Biochim. Biophys. Acta / Probing the structure of Lhca3 by mutation analysis by Mozzo (2006)
10.1529/biophysj.107.106955 / Biophys. J. / The low-energy forms of Photosystem I light-harvesting complexes: spectroscopic properties and pigment–pigment interaction characteristics by Croce (2007)
10.1021/jp0519316 / J. Phys. Chem. B / Excitation decay pathways of Lhca proteins: a time-resolved fluorescence study by Ihalainen (2005)
10.1021/jp034778t / J. Phys. Chem. B / Red spectral forms of chlorophylls in green plant PSI: a site-selective and high-pressure spectroscopy study by Ihalainen (2003)
10.1016/j.bpj.2008.11.043 / Biophys. J. / The origin of the low-energy form of Photosystem I light-harvesting complex Lhca4: mixing of the lowest exciton with a charge-transfer state by Romero (2009)
10.1104/pp.103.033324 / Plant Physiol. / Is each light-harvesting complex protein important for plant fitness? by Ganeteg (2004)
10.1104/pp.127.1.150 / Plant Physiol. / The properties of the chlorophyll a/b-binding proteins Lhca2 and Lhca3 studied in vivo using antisense inhibition by Ganeteg (2001)
10.1074/jbc.M808395200 / J. Biol. Chem. / The role of Lhca complexes in the supramolecular organization of higher plant Photosystem I by Wientjes (2009)
10.1021/bi0257437 / Biochemistry / Chromophore organization in the higher-plant photosystem II antenna protein CP26 by Croce (2002)
10.1039/tf9575300646 / Trans. Faraday Soc. / Determination of the absolute quantum yield of fluorescent solutions by Weber (1957)
10.1021/bi047873g / Biochemistry / Structure of the higher plant light harvesting complex I: in vivo characterization and structural interdependence of the Lhca proteins by Klimmek (2005)
10.1016/j.bbabio.2008.04.036 / Biochim. Biophys. Acta / Photoprotection in higher plants: the putative quenching site is conserved in all outer light-harvesting complexes of Photosystem II by Mozzo (2008)
10.1146/annurev.arplant.47.1.655 / Annu. Rev. Plant Physiol. Plant Mol. Biol. / Regulation of light harvesting in green plants by Horton (1996)
10.1021/jp014078t / J. Phys. Chem. B / Superradiance and exciton (de)localization in light-harvesting complex II from green plants? by Palacios (2002)
10.1142/3609 / Photosynthetic Excitons by van Amerongen (2000)
10.1021/bi050260z / Biochemistry / Quenching of chlorophyll triplet states by carotenoids in reconstituted Lhca4 subunit of peripheral light-harvesting complex of Photosystem I by Carbonera (2005)
10.1021/bi026157g / Biochemistry / Light-induced trimer to monomer transition in the main light-harvesting antenna complex of plants: thermo-optic mechanism by Garab (2002)
10.1007/978-1-4020-4256-0_10 / Photosystem I: The Light-Driven Plastocyanin: Ferredoxin Oxidoreductase / LHCI: the antenna complex of Photosystem I in plants and green algae by Croce (2006)
10.1074/jbc.M502935200 / J. Biol. Chem. / The association of the antenna system to Photosystem I in higher plants by Morosinotto (2005)
10.1021/bi602531k / Biochemistry / Singlet and triplet state transitions of carotenoids in the antenna complexes of higher-plant Photosystem I by Croce (2007)
10.1111/j.1742-4658.2004.04512.x / FEBS J. / Protective effect of active oxygen scavengers on protein degradation and photochemical function in photosystem I submembrane fractions during light stress by Rajagopal (2005)
10.1007/BF02900557 / Chin. Sci. Bull. / The photodamage process of pigments and proteins of PSI complexes from Spinacia oleracea L by Wei (2001)

Dates

Type	When
Created	14 years, 9 months ago (Nov. 19, 2010, 10:52 a.m.)
Deposited	3 years, 9 months ago (Nov. 22, 2021, 2:07 p.m.)
Indexed	1 week, 1 day ago (Aug. 29, 2025, 5:48 a.m.)
Issued	14 years, 7 months ago (Jan. 14, 2011)
Published	14 years, 7 months ago (Jan. 14, 2011)
Published Online	14 years, 7 months ago (Jan. 14, 2011)
Published Print	14 years, 7 months ago (Feb. 1, 2011)

Funders 0

None

BibTeX

@article{Wientjes_2011, title={The light-harvesting complexes of higher-plant Photosystem I: Lhca1/4 and Lhca2/3 form two red-emitting heterodimers}, volume={433}, ISSN={1470-8728}, url={http://dx.doi.org/10.1042/bj20101538}, DOI={10.1042/bj20101538}, number={3}, journal={Biochemical Journal}, publisher={Portland Press Ltd.}, author={Wientjes, Emilie and Croce, Roberta}, year={2011}, month=jan, pages={477–485} }

JSON

{
  "indexed": {
    "date-parts": [
      [
        2025,
        8,
        29
      ]
    ],
    "date-time": "2025-08-29T09:48:10Z",
    "timestamp": 1756460890879
  },
  "reference-count": 56,
  "publisher": "Portland Press Ltd.",
  "issue": "3",
  "content-domain": {
    "domain": [
      "portlandpress.com"
    ],
    "crossmark-restriction": true
  },
  "published-print": {
    "date-parts": [
      [
        2011,
        2,
        1
      ]
    ]
  },
  "abstract": "<jats:p>The outer antenna of higher-plant PSI (Photosystem I) is composed of four complexes [Lhc (light-harvesting complex) a1\u2013Lhca4] belonging to the light-harvesting protein family. Difficulties in their purification have so far prevented the determination of their properties and most of the knowledge about Lhcas has been obtained from the study of the in vitro reconstituted antennas. In the present study we were able to purify the native complexes, showing that Lhca2/3 and Lhca1/4 form two functional heterodimers. Both dimers show red-fluorescence emission with maxima around 730\u00a0nm, as in the intact PSI complex. This indicates that the dimers are in their native state and that LHCI-680, which was previously assumed to be part of the PSI antenna, does not represent the native state of the system. The data show that the light-harvesting properties of the two dimers are functionally identical, concerning absorption, long-wavelength emission and fluorescence quantum yield, whereas they differ in their high-light response. Implications of the present study for the understanding of the energy transfer process in PSI are discussed. Finally, the comparison of the properties of the native dimers with those of the reconstituted complexes demonstrates that all of the major properties of the Lhcas are reproduced in the in vitro systems.</jats:p>",
  "DOI": "10.1042/bj20101538",
  "type": "journal-article",
  "created": {
    "date-parts": [
      [
        2010,
        11,
        19
      ]
    ],
    "date-time": "2010-11-19T15:52:33Z",
    "timestamp": 1290181953000
  },
  "page": "477-485",
  "update-policy": "http://dx.doi.org/10.1042/crossmark_policy",
  "source": "Crossref",
  "is-referenced-by-count": 105,
  "title": "The light-harvesting complexes of higher-plant Photosystem I: Lhca1/4 and Lhca2/3 form two red-emitting heterodimers",
  "prefix": "10.1042",
  "volume": "433",
  "author": [
    {
      "given": "Emilie",
      "family": "Wientjes",
      "sequence": "first",
      "affiliation": [
        {
          "name": "Department of Biophysical Chemistry, Groningen Biomolecular Sciences and Biotechnology Institute, University of Groningen, Nijenborgh 4, 9747 AG Groningen, The Netherlands"
        }
      ]
    },
    {
      "given": "Roberta",
      "family": "Croce",
      "sequence": "additional",
      "affiliation": [
        {
          "name": "Department of Biophysical Chemistry, Groningen Biomolecular Sciences and Biotechnology Institute, University of Groningen, Nijenborgh 4, 9747 AG Groningen, The Netherlands"
        }
      ]
    }
  ],
  "member": "288",
  "published-online": {
    "date-parts": [
      [
        2011,
        1,
        14
      ]
    ]
  },
  "reference": [
    {
      "key": "2021112214452763100_B1",
      "doi-asserted-by": "crossref",
      "first-page": "58",
      "DOI": "10.1038/nature05687",
      "article-title": "The structure of a plant Photosystem I supercomplex at 3.4 \u00c5 resolution",
      "volume": "447",
      "author": "Amunts",
      "year": "2007",
      "journal-title": "Nature"
    },
    {
      "key": "2021112214452763100_B2",
      "doi-asserted-by": "crossref",
      "first-page": "909",
      "DOI": "10.1038/35082000",
      "article-title": "Three-dimensional structure of cyanobacterial Photosystem I at 2.5 \u00c5 resolution",
      "volume": "411",
      "author": "Jordan",
      "year": "2001",
      "journal-title": "Nature"
    },
    {
      "key": "2021112214452763100_B3",
      "doi-asserted-by": "crossref",
      "first-page": "421",
      "DOI": "10.1007/978-94-011-3953-3_100",
      "article-title": "The light-harvesting complex of photosytem I: pigment composition and stoichiometry",
      "volume-title": "Photosynthesis: Mechanisms and Effects",
      "author": "Croce",
      "year": "1998"
    },
    {
      "key": "2021112214452763100_B4",
      "doi-asserted-by": "crossref",
      "first-page": "236",
      "DOI": "10.1016/S1360-1385(99)01419-3",
      "article-title": "A guide to the Lhc genes and their relatives in Arabidopsis",
      "volume": "4",
      "author": "Jansson",
      "year": "1999",
      "journal-title": "Trends Plant Sci."
    },
    {
      "key": "2021112214452763100_B5",
      "doi-asserted-by": "crossref",
      "first-page": "1029",
      "DOI": "10.1021/bi0015358",
      "article-title": "Green plant Photosystem I binds light-harvesting complex I on one side of the complex",
      "volume": "40",
      "author": "Boekema",
      "year": "2001",
      "journal-title": "Biochemistry"
    },
    {
      "key": "2021112214452763100_B6",
      "doi-asserted-by": "crossref",
      "first-page": "630",
      "DOI": "10.1038/nature02200",
      "article-title": "Crystal structure of plant Photosystem I",
      "volume": "426",
      "author": "Ben-Shem",
      "year": "2003",
      "journal-title": "Nature"
    },
    {
      "key": "2021112214452763100_B7",
      "doi-asserted-by": "crossref",
      "first-page": "4659",
      "DOI": "10.1111/j.1432-1033.2004.04426.x",
      "article-title": "Stoichiometry of LHCI antenna polypeptides and characterization of gap and linker pigments in higher plants Photosystem I",
      "volume": "271",
      "author": "Ballottari",
      "year": "2004",
      "journal-title": "Eur. J. Biochem."
    },
    {
      "key": "2021112214452763100_B8",
      "doi-asserted-by": "crossref",
      "first-page": "8947",
      "DOI": "10.1074/jbc.M606417200",
      "article-title": "Contrasting behavior of higher plant Photosystem I and II antenna systems during acclimation",
      "volume": "282",
      "author": "Ballottari",
      "year": "2007",
      "journal-title": "J. Biol. Chem."
    },
    {
      "key": "2021112214452763100_B9",
      "doi-asserted-by": "crossref",
      "first-page": "71",
      "DOI": "10.1186/1471-2229-9-71",
      "article-title": "Antenna complexes protect Photosystem I from photoinhibition",
      "volume": "9",
      "author": "Alboresi",
      "year": "2009",
      "journal-title": "BMC Plant Biol."
    },
    {
      "key": "2021112214452763100_B10",
      "doi-asserted-by": "crossref",
      "first-page": "1301",
      "DOI": "10.1111/j.1751-1097.2007.00136.x",
      "article-title": "Selective photobleaching of chlorophylls and carotenoids in Photosystem I particles under high-light treatment",
      "volume": "83",
      "author": "Andreeva",
      "year": "2007",
      "journal-title": "Photochem. Photobiol."
    },
    {
      "key": "2021112214452763100_B11",
      "doi-asserted-by": "crossref",
      "first-page": "508",
      "DOI": "10.1562/0031-8655(2000)072<0508:DOTPIC>2.0.CO;2",
      "article-title": "Degradation of the Photosystem I complex during photoinhibition",
      "volume": "72",
      "author": "Hui",
      "year": "2000",
      "journal-title": "Photochem. Photobiol."
    },
    {
      "key": "2021112214452763100_B12",
      "doi-asserted-by": "crossref",
      "first-page": "80",
      "DOI": "10.1016/S0005-2728(01)00203-1",
      "article-title": "Energy transfer and trapping in Photosystem I",
      "volume": "1507",
      "author": "Gobets",
      "year": "2001",
      "journal-title": "Biochim. Biophys. Acta"
    },
    {
      "key": "2021112214452763100_B13",
      "doi-asserted-by": "crossref",
      "first-page": "209",
      "DOI": "10.1023/A:1006236829711",
      "article-title": "The importance of PSI chlorophyll red forms in light-harvesting by leaves",
      "volume": "60",
      "author": "Rivadossi",
      "year": "1999",
      "journal-title": "Photosyn. Res."
    },
    {
      "key": "2021112214452763100_B14",
      "doi-asserted-by": "crossref",
      "first-page": "247",
      "DOI": "10.1007/BF00016556",
      "article-title": "Long-wavelength absorbing antenna pigments and heterogeneous absorption-bands concentrate excitons and increase absorption cross-section",
      "volume": "35",
      "author": "Trissl",
      "year": "1993",
      "journal-title": "Photosyn. Res."
    },
    {
      "key": "2021112214452763100_B15",
      "doi-asserted-by": "crossref",
      "first-page": "267",
      "DOI": "10.1016/j.bbabio.2004.11.007",
      "article-title": "Kinetics of excitation trapping in intact Photosystem I of Chlamydomonas reinhardtii and Arabidopsis thaliana",
      "volume": "1706",
      "author": "Ihalainen",
      "year": "2005",
      "journal-title": "Biochim. Biophys. Acta"
    },
    {
      "key": "2021112214452763100_B16",
      "doi-asserted-by": "crossref",
      "first-page": "91",
      "DOI": "10.1016/S0005-2728(02)00399-7",
      "article-title": "The photochemical trapping rate from red spectral states in PSI-LHCI is determined by thermal activation of energy transfer to bulk chlorophylls",
      "volume": "1557",
      "author": "Jennings",
      "year": "2003",
      "journal-title": "Biochim. Biophys. Acta"
    },
    {
      "key": "2021112214452763100_B17",
      "doi-asserted-by": "crossref",
      "first-page": "8572",
      "DOI": "10.1021/bi960214m",
      "article-title": "Excited state equilibration in the Photosystem I light-harvesting I complex: P700 is almost isoenergetic with its antenna",
      "volume": "35",
      "author": "Croce",
      "year": "1996",
      "journal-title": "Biochemistry"
    },
    {
      "key": "2021112214452763100_B18",
      "doi-asserted-by": "crossref",
      "first-page": "3601",
      "DOI": "10.1529/biophysj.107.117101",
      "article-title": "Trap-limited charge separation kinetics in higher plant Photosystem I complexes",
      "volume": "94",
      "author": "Slavov",
      "year": "2008",
      "journal-title": "Biophys. J."
    },
    {
      "key": "2021112214452763100_B19",
      "doi-asserted-by": "crossref",
      "first-page": "5851",
      "DOI": "10.1529/biophysj.108.140467",
      "article-title": "Picosecond fluorescence of intact and dissolved PSI-LHCI crystals",
      "volume": "95",
      "author": "van\u00a0Oort",
      "year": "2008",
      "journal-title": "Biophys. J."
    },
    {
      "key": "2021112214452763100_B20",
      "doi-asserted-by": "crossref",
      "first-page": "221",
      "DOI": "10.1111/j.1432-1033.1987.tb10791.x",
      "article-title": "Chlorophyll\u2013protein complexes of barley Photosystem-I",
      "volume": "163",
      "author": "Bassi",
      "year": "1987",
      "journal-title": "Eur. J. Biochem."
    },
    {
      "key": "2021112214452763100_B21",
      "doi-asserted-by": "crossref",
      "first-page": "209",
      "DOI": "10.1111/j.1432-1033.1992.tb16918.x",
      "article-title": "Identification of the Photosystem I antenna polypeptides in barley: isolation of three pigment-binding antenna complexes",
      "volume": "206",
      "author": "Knoetzel",
      "year": "1992",
      "journal-title": "Eur. J. Biochem."
    },
    {
      "key": "2021112214452763100_B22",
      "doi-asserted-by": "crossref",
      "first-page": "10",
      "DOI": "10.1016/0014-5793(84)80197-0",
      "article-title": "Chlorophyll a/b proteins of Photosystem I",
      "volume": "168",
      "author": "Lam",
      "year": "1984",
      "journal-title": "FEBS Lett."
    },
    {
      "key": "2021112214452763100_B23",
      "doi-asserted-by": "crossref",
      "first-page": "37307",
      "DOI": "10.1074/jbc.M205889200",
      "article-title": "Pigment binding of Photosystem I light-harvesting proteins",
      "volume": "277",
      "author": "Schmid",
      "year": "2002",
      "journal-title": "J. Biol. Chem."
    },
    {
      "key": "2021112214452763100_B24",
      "doi-asserted-by": "crossref",
      "first-page": "41",
      "DOI": "10.1007/BF00032234",
      "article-title": "Rapid isolation of Photosystem I chlorophyll-binding proteins by anion-exchange perfusion chromatography",
      "volume": "45",
      "author": "Tjus",
      "year": "1995",
      "journal-title": "Photosyn. Res."
    },
    {
      "key": "2021112214452763100_B25",
      "doi-asserted-by": "crossref",
      "first-page": "17355",
      "DOI": "10.1021/bi9813227",
      "article-title": "A thermal broadening study of the antenna chlorophylls in PSI-200, LHCI, and PSI core",
      "volume": "37",
      "author": "Croce",
      "year": "1998",
      "journal-title": "Biochemistry"
    },
    {
      "key": "2021112214452763100_B26",
      "doi-asserted-by": "crossref",
      "first-page": "8625",
      "DOI": "10.1021/bi0007369",
      "article-title": "Evidence for two spectroscopically different dimers of light-harvesting complex I from green plants",
      "volume": "39",
      "author": "Ihalainen",
      "year": "2000",
      "journal-title": "Biochemistry"
    },
    {
      "key": "2021112214452763100_B27",
      "first-page": "29",
      "article-title": "The Lhca antenna complexes of higher plants photosystem I. Biochim",
      "volume": "1556",
      "author": "Croce",
      "year": "2002",
      "journal-title": "Biophy. Acta"
    },
    {
      "key": "2021112214452763100_B28",
      "doi-asserted-by": "crossref",
      "first-page": "4226",
      "DOI": "10.1021/bi027398r",
      "article-title": "Recombinant Lhca2 and Lhca3 subunits of the Photosystem I antenna system",
      "volume": "42",
      "author": "Castelletti",
      "year": "2003",
      "journal-title": "Biochemistry"
    },
    {
      "key": "2021112214452763100_B29",
      "doi-asserted-by": "crossref",
      "first-page": "7667",
      "DOI": "10.1073/pnas.94.14.7667",
      "article-title": "In vitro reconstitution of the Photosystem I light-harvesting complex LHCI-730: heterodimerization is required for antenna pigment organization",
      "volume": "94",
      "author": "Schmid",
      "year": "1997",
      "journal-title": "Proc. Natl. Acad. Sci. U.S.A."
    },
    {
      "key": "2021112214452763100_B30",
      "doi-asserted-by": "crossref",
      "first-page": "287",
      "DOI": "10.1038/nature02373",
      "article-title": "Crystal structure of spinach major light-harvesting complex at 2.72 \u00c5 resolution",
      "volume": "428",
      "author": "Liu",
      "year": "2004",
      "journal-title": "Nature"
    },
    {
      "key": "2021112214452763100_B31",
      "doi-asserted-by": "crossref",
      "first-page": "48543",
      "DOI": "10.1074/jbc.M408908200",
      "article-title": "Origin of the 701\u00a0nm fluorescence emission of the Lhca2 subunit of higher plant Photosystem I",
      "volume": "279",
      "author": "Croce",
      "year": "2004",
      "journal-title": "J. Biol. Chem."
    },
    {
      "key": "2021112214452763100_B32",
      "doi-asserted-by": "crossref",
      "first-page": "49223",
      "DOI": "10.1074/jbc.M309203200",
      "article-title": "The nature of a chlorophyll ligand in Lhca proteins determines the far red fluorescence emission typical of Photosystem I",
      "volume": "278",
      "author": "Morosinotto",
      "year": "2003",
      "journal-title": "J. Biol. Chem."
    },
    {
      "key": "2021112214452763100_B33",
      "doi-asserted-by": "crossref",
      "first-page": "36253",
      "DOI": "10.1074/jbc.M205062200",
      "article-title": "Mutation analysis of Lhca1 antenna complex: low energy absorption forms originate from pigment\u2013pigment interactions",
      "volume": "277",
      "author": "Morosinotto",
      "year": "2002",
      "journal-title": "J. Biol. Chem."
    },
    {
      "key": "2021112214452763100_B34",
      "doi-asserted-by": "crossref",
      "first-page": "20612",
      "DOI": "10.1074/jbc.M500705200",
      "article-title": "Pigment-pigment interactions in Lhca4 antenna complex of higher plants Photosystem I",
      "volume": "280",
      "author": "Morosinotto",
      "year": "2005",
      "journal-title": "J. Biol. Chem."
    },
    {
      "key": "2021112214452763100_B35",
      "doi-asserted-by": "crossref",
      "first-page": "1607",
      "DOI": "10.1016/j.bbabio.2006.06.018",
      "article-title": "Probing the structure of Lhca3 by mutation analysis",
      "volume": "1757",
      "author": "Mozzo",
      "year": "2006",
      "journal-title": "Biochim. Biophys. Acta"
    },
    {
      "key": "2021112214452763100_B36",
      "doi-asserted-by": "crossref",
      "first-page": "2418",
      "DOI": "10.1529/biophysj.107.106955",
      "article-title": "The low-energy forms of Photosystem I light-harvesting complexes: spectroscopic properties and pigment\u2013pigment interaction characteristics",
      "volume": "93",
      "author": "Croce",
      "year": "2007",
      "journal-title": "Biophys. J."
    },
    {
      "key": "2021112214452763100_B37",
      "doi-asserted-by": "crossref",
      "first-page": "21150",
      "DOI": "10.1021/jp0519316",
      "article-title": "Excitation decay pathways of Lhca proteins: a time-resolved fluorescence study",
      "volume": "109",
      "author": "Ihalainen",
      "year": "2005",
      "journal-title": "J. Phys. Chem. B"
    },
    {
      "key": "2021112214452763100_B38",
      "doi-asserted-by": "crossref",
      "first-page": "9086",
      "DOI": "10.1021/jp034778t",
      "article-title": "Red spectral forms of chlorophylls in green plant PSI: a site-selective and high-pressure spectroscopy study",
      "volume": "107",
      "author": "Ihalainen",
      "year": "2003",
      "journal-title": "J. Phys. Chem. B"
    },
    {
      "key": "2021112214452763100_B39",
      "doi-asserted-by": "crossref",
      "first-page": "L35",
      "DOI": "10.1016/j.bpj.2008.11.043",
      "article-title": "The origin of the low-energy form of Photosystem I light-harvesting complex Lhca4: mixing of the lowest exciton with a charge-transfer state",
      "volume": "96",
      "author": "Romero",
      "year": "2009",
      "journal-title": "Biophys. J."
    },
    {
      "key": "2021112214452763100_B40",
      "doi-asserted-by": "crossref",
      "first-page": "502",
      "DOI": "10.1104/pp.103.033324",
      "article-title": "Is each light-harvesting complex protein important for plant fitness?",
      "volume": "134",
      "author": "Ganeteg",
      "year": "2004",
      "journal-title": "Plant Physiol."
    },
    {
      "key": "2021112214452763100_B41",
      "doi-asserted-by": "crossref",
      "first-page": "150",
      "DOI": "10.1104/pp.127.1.150",
      "article-title": "The properties of the chlorophyll a/b-binding proteins Lhca2 and Lhca3 studied in vivo using antisense inhibition",
      "volume": "127",
      "author": "Ganeteg",
      "year": "2001",
      "journal-title": "Plant Physiol."
    },
    {
      "key": "2021112214452763100_B42",
      "doi-asserted-by": "crossref",
      "first-page": "7803",
      "DOI": "10.1074/jbc.M808395200",
      "article-title": "The role of Lhca complexes in the supramolecular organization of higher plant Photosystem I",
      "volume": "284",
      "author": "Wientjes",
      "year": "2009",
      "journal-title": "J. Biol. Chem."
    },
    {
      "key": "2021112214452763100_B43",
      "doi-asserted-by": "crossref",
      "first-page": "7334",
      "DOI": "10.1021/bi0257437",
      "article-title": "Chromophore organization in the higher-plant photosystem II antenna protein CP26",
      "volume": "41",
      "author": "Croce",
      "year": "2002",
      "journal-title": "Biochemistry"
    },
    {
      "key": "2021112214452763100_B44",
      "doi-asserted-by": "crossref",
      "first-page": "646",
      "DOI": "10.1039/tf9575300646",
      "article-title": "Determination of the absolute quantum yield of fluorescent solutions",
      "volume": "53",
      "author": "Weber",
      "year": "1957",
      "journal-title": "Trans. Faraday Soc."
    },
    {
      "key": "2021112214452763100_B45",
      "doi-asserted-by": "crossref",
      "first-page": "3065",
      "DOI": "10.1021/bi047873g",
      "article-title": "Structure of the higher plant light harvesting complex I: in vivo characterization and structural interdependence of the Lhca proteins",
      "volume": "44",
      "author": "Klimmek",
      "year": "2005",
      "journal-title": "Biochemistry"
    },
    {
      "key": "2021112214452763100_B46",
      "doi-asserted-by": "crossref",
      "first-page": "1263",
      "DOI": "10.1016/j.bbabio.2008.04.036",
      "article-title": "Photoprotection in higher plants: the putative quenching site is conserved in all outer light-harvesting complexes of Photosystem II",
      "volume": "1777",
      "author": "Mozzo",
      "year": "2008",
      "journal-title": "Biochim. Biophys. Acta"
    },
    {
      "key": "2021112214452763100_B47",
      "doi-asserted-by": "crossref",
      "first-page": "655",
      "DOI": "10.1146/annurev.arplant.47.1.655",
      "article-title": "Regulation of light harvesting in green plants",
      "volume": "47",
      "author": "Horton",
      "year": "1996",
      "journal-title": "Annu. Rev. Plant Physiol. Plant Mol. Biol."
    },
    {
      "key": "2021112214452763100_B48",
      "doi-asserted-by": "crossref",
      "first-page": "5782",
      "DOI": "10.1021/jp014078t",
      "article-title": "Superradiance and exciton (de)localization in light-harvesting complex II from green plants?",
      "volume": "106",
      "author": "Palacios",
      "year": "2002",
      "journal-title": "J. Phys. Chem. B"
    },
    {
      "key": "2021112214452763100_B49",
      "doi-asserted-by": "crossref",
      "DOI": "10.1142/3609",
      "volume-title": "Photosynthetic Excitons",
      "author": "van\u00a0Amerongen",
      "year": "2000"
    },
    {
      "key": "2021112214452763100_B50",
      "doi-asserted-by": "crossref",
      "first-page": "8337",
      "DOI": "10.1021/bi050260z",
      "article-title": "Quenching of chlorophyll triplet states by carotenoids in reconstituted Lhca4 subunit of peripheral light-harvesting complex of Photosystem I",
      "volume": "44",
      "author": "Carbonera",
      "year": "2005",
      "journal-title": "Biochemistry"
    },
    {
      "key": "2021112214452763100_B51",
      "doi-asserted-by": "crossref",
      "first-page": "15121",
      "DOI": "10.1021/bi026157g",
      "article-title": "Light-induced trimer to monomer transition in the main light-harvesting antenna complex of plants: thermo-optic mechanism",
      "volume": "41",
      "author": "Garab",
      "year": "2002",
      "journal-title": "Biochemistry"
    },
    {
      "key": "2021112214452763100_B52",
      "doi-asserted-by": "crossref",
      "first-page": "119",
      "DOI": "10.1007/978-1-4020-4256-0_10",
      "article-title": "LHCI: the antenna complex of Photosystem I in plants and green algae",
      "volume-title": "Photosystem I: The Light-Driven Plastocyanin: Ferredoxin Oxidoreductase",
      "author": "Croce",
      "year": "2006"
    },
    {
      "key": "2021112214452763100_B53",
      "doi-asserted-by": "crossref",
      "first-page": "31050",
      "DOI": "10.1074/jbc.M502935200",
      "article-title": "The association of the antenna system to Photosystem I in higher plants",
      "volume": "280",
      "author": "Morosinotto",
      "year": "2005",
      "journal-title": "J. Biol. Chem."
    },
    {
      "key": "2021112214452763100_B54",
      "doi-asserted-by": "crossref",
      "first-page": "3846",
      "DOI": "10.1021/bi602531k",
      "article-title": "Singlet and triplet state transitions of carotenoids in the antenna complexes of higher-plant Photosystem I",
      "volume": "46",
      "author": "Croce",
      "year": "2007",
      "journal-title": "Biochemistry"
    },
    {
      "key": "2021112214452763100_B55",
      "doi-asserted-by": "crossref",
      "first-page": "892",
      "DOI": "10.1111/j.1742-4658.2004.04512.x",
      "article-title": "Protective effect of active oxygen scavengers on protein degradation and photochemical function in photosystem I submembrane fractions during light stress",
      "volume": "272",
      "author": "Rajagopal",
      "year": "2005",
      "journal-title": "FEBS J."
    },
    {
      "key": "2021112214452763100_B56",
      "doi-asserted-by": "crossref",
      "first-page": "1812",
      "DOI": "10.1007/BF02900557",
      "article-title": "The photodamage process of pigments and proteins of PSI complexes from Spinacia oleracea L",
      "volume": "46",
      "author": "Wei",
      "year": "2001",
      "journal-title": "Chin. Sci. Bull."
    }
  ],
  "container-title": "Biochemical Journal",
  "original-title": [],
  "language": "en",
  "link": [
    {
      "URL": "https://portlandpress.com/biochemj/article-pdf/433/3/477/664693/bj4330477.pdf",
      "content-type": "application/pdf",
      "content-version": "vor",
      "intended-application": "syndication"
    },
    {
      "URL": "https://portlandpress.com/biochemj/article-pdf/433/3/477/664693/bj4330477.pdf",
      "content-type": "unspecified",
      "content-version": "vor",
      "intended-application": "similarity-checking"
    }
  ],
  "deposited": {
    "date-parts": [
      [
        2021,
        11,
        22
      ]
    ],
    "date-time": "2021-11-22T19:07:04Z",
    "timestamp": 1637608024000
  },
  "score": 1,
  "resource": {
    "primary": {
      "URL": "https://portlandpress.com/biochemj/article/433/3/477/45486/The-light-harvesting-complexes-of-higher-plant"
    }
  },
  "subtitle": [],
  "short-title": [],
  "issued": {
    "date-parts": [
      [
        2011,
        1,
        14
      ]
    ]
  },
  "references-count": 56,
  "journal-issue": {
    "issue": "3",
    "published-print": {
      "date-parts": [
        [
          2011,
          2,
          1
        ]
      ]
    }
  },
  "URL": "http://dx.doi.org/10.1042/BJ20101538",
  "relation": {},
  "ISSN": [
    "0264-6021",
    "1470-8728"
  ],
  "subject": [],
  "published": {
    "date-parts": [
      [
        2011,
        1,
        14
      ]
    ]
  }
}