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Photosynthesis and the Web: 2008 Subject SitesBroad Overview SitesThere are several sites devoted to the entire photosynthetic process. Although they may cover the same subject, the articles are written for a variety of audiences. Some are basic narratives aimed at the general public, others are comprehensive courses aimed at college students. All of these are quite good and several should be checked out for comparison. "The Photosynthetic Process" by John Whitmarsh and Govindjee (http://www.life.uiuc.edu/govindjee/paper/gov.html) is a comprehensive chapter and covers photosynthesis history and every facet of photosynthesis research in a relatively detailed manner. Wim Vermaas at ASU has written "An Introduction to Photosynthesis and Its Applications" (http://photoscience.la.asu.edu/photosyn/education/photointro.html) which is a good introduction for the general public. J. M Farabee from Estrella Mountain Community College also has written a highly recommended online biology course text that includes a detailed section on photosynthesis with emphasis on the physical aspects of the process. He also includes review questions, learning objectives and a nicely done illustrated glossary (http://www.emc.maricopa.edu/faculty/farabee/BIOBK/BioBookPS.html). Members of the Faculty of Biology at the University of Hamburg, Germany have also produced an online botany textbook with excellent section devoted to photosynthesis (http://www.biologie.uni-hamburg.de/b-online/e24/24.htm). Kapiolani Community College in Hawaii has a web site devoted to the chemical equations of photosynthesis (http://library.kcc.hawaii.edu/external/chemistry/everyday_photosyn.html). Although the online Encyclopaedia Britannica is only available by subscription, the entire 36 page section on photosynthesis is available at no cost through a link on the Photosynthesis Center educational resources page as the Center is a recognized Britannica iGuide site (http://photoscience.la.asu.edu/photosyn/education.html). Wikipedia has extensive coverage of the entire process with many links at http://en.wikipedia.org/wiki/Photosynthesis. Both authors of this review have participated in the editing of the Wikipedia. Please note that the Wikipedia is a work in progress and is far from perfect. Because of its open communal editing process, not all information is entirely accurate and much more space is devoted to items of interest to those that edit the Wikipedia, i.e., current popular culture such as rock groups, movie stars, etc., than science. Still, it is a good place to look for quick information even on scientific subjects, though it is recommended that another source always be checked before relying completely on its contents for one’s research. If you have the time, we encourage everyone to open a no-charge editorial account and help maintain this interesting venture. June B. Steinberg's site at http://faculty.nl.edu/jste/photosynthesis.htm has easy-to-understand explanations and links to more detailed information. There are many useful animations . Molecular Expressions at Florida State University has a wonderful animated web page as part of its Optical Microscopy Primer (http://micro.magnet.fsu.edu/primer/java/photosynthesis/index.html). There is much here to enjoy and it is fun to roam around through the site. For example, see the section on "Light and Color" (http://micro.magnet.fsu.edu/primer/lightandcolor/index.html) with defines and explains what light is and concepts such as fluorescence, reflection, refraction, human perception of color, lasers, etc. A truly wonderful sight. FT Exploring has a wonderful site, "PHOTOSYNTHESIS: HOW LIFE KEEPS GOING...and going...and going...," for students (http://www.ftexploring.com/photosyn/photosynth.html). It covers the details of photosynthesis and how it relates to the rest of life. There is much good information here, as well as good humor and illustrations. Specific Subject SitesThere are many sites that specialize in a certain aspect of photosynthesis. Here are some of the best arranged by topic. There will be several sites that overlap somewhat and some may appear in more than one group. History of Photosynthesis/Biographies/Nobel Prizes"Milestones in Photosynthesis Research" by one of us (G) (http://www.life.uiuc.edu/govindjee/papers/milestones.html) explores many aspects of photosynthesis in a historic manner. Another paper, "Carotenoids in Photosynthesis: An Historical Perspective," explores the history of the study of carotenoids using numerous personal observations by one of the authors (G) who participated in many of the activities (http://www.life.uiuc.edu/govindjee/papers/CarFin1.html). Both of these papers contain numerous references and anecdotes about pioneers in the field that can be found nowhere else. A list of historical articles, published in Photosynthesis Research, are available at: http://www.life.uiuc.edu/govindjee/history/articles.htm. Further, PDF files of articles by Howard Gest (on IngenHousz); Herb Dutton (on the discovery of energy transfer from carotenoids to chlorophyll); and Govindjee (on the quantum yield controversy between Emerson and Warburg) are also available at this site. Other important historical documents are: "Nobel Prize Winners in Photosynthesis Research" (http://www.life.uiuc.edu/govindjee/history/nobel-ps.htm) is a list of the many researchers who have won the Nobel Prize while or before studying photosynthesis. "Personal Perspectives in Photosynthesis Research" (http://www.life.uiuc.edu/govindjee/perspectives.html) is a list of the Personal Perspectives, autobiographical retrospectives, that have appeared in the international journal Photosynthesis Research. Although these perspectives usually do not contain large amounts of science, they do discuss the major discoveries by some well-known researchers. In particular, they contain the personal reflections, memories and the obstacles the authors had to overcome and other surprises. For example, the Personal Perspective of David Walker begins, "This is the story of a young man who wished to go to sea like his father and finished up, instead, in photosynthesis." They are great reading and tell about scientific research the way it really is. 111 of these perspectives by 132 authors in 19 countries have been collected in a recent book, Discoveries in Photosynthesis, edited by Govindjee, J. Thomas Beatty, Howard Gest and John F. Allen (http://www.life.uiuc.edu/govindjee/newbook/Vol%2020.html). A few of the perspectives are available as PDF files (please note that the first page of each file may be blank):
Govindjee has also edited several obituaries that have been published (http://www.life.uiuc.edu/govindjee/history/obituaries.htm), with some of them available online as PDF files. Carmen Giunta has collected excerpts from historically important papers and published them on his "Classic Chemistry" web site. These include papers by Jan Ingenhousz (http://webserver.lemoyne.edu/faculty/giunta/Ingenhousz.html), Antoine Lavoisier (http://webserver.lemoyne.edu/faculty/giunta/lavoisier1.html), Joseph Priestley (http://webserver.lemoyne.edu/faculty/giunta/phlogiston.html). A brief biography of Joseph Priestley can be found at http://www.chemistry.mtu.edu/~pcharles/SCIHISTORY/JosephPriestley.html. A brief biographies of Jan Ingenhousz can be found at http://www.chemheritage.org/explore/life-ingenhousz.html and http://www.newadvent.org/cathen/16046b.htm The Nobel Prize site has pages devoted to all Nobel laureates. Some of interest to photosynthesis are:
Photograph of Richard Martin Willstatter Richard Martin Willstatter, Chemistry, 1915, won the prize for his research on chlorophyll and other plant pigments. His work is discussed at: http://www.nobel.se/chemistry/laureates/1915/press.html and his biography is at: http://www.nobel.se/chemistry/laureates/1915/willstatter-bio.html James Franck, Physics, 1925, won for his work (with Gustav Hertz) on electron-atom collisions; later he developed the principle known as the Franck-Condon principle which is often used in physical description of early events in photosynthesis. See (http://www.nobel.se/physics/laureates/1925/press.html) and his biography (http://www.nobel.se/physics/laureates/1925/franck-bio.html) Chandrasekhara Ventaka Raman, Physics, 1930, won his prize for his work on spectroscopy and the effect that now bears his name, Raman spectroscopy, which is used by many photosynthesis researchers: (http://www.nobel.se/physics/laureates/1930/press.html). His biography is at (http://www.nobel.se/physics/laureates/1930/raman-bio.html). Hans Fischer, Chemistry, 1930, won for his work on porphyrins and blood and leaf pigments, particularly chlorophyll: http://www.nobel.se/chemistry/laureates/1930/press.html. His biography is at (http://www.nobel.se/chemistry/laureates/1930/fischer-bio.html) Otto Heinrich Warburg, Physiology or Medicine, 1931, won for his work on respiration and the identification of the respiratory enzyme: http://www.nobel.se/medicine/laureates/1931/press.html. His biography is at (http://www.nobel.se/medicine/laureates/1931/warburg-bio.html). Warburg's insistence that the measured minimum quantum requirement for the evolution of one oxygen molecule in photosynthesis is 2.8 to 4 was proven to be wrong; it was shown to be 8-12, mainly by Robert Emerson and his students including one of us (G). Paul Karrer, Chemistry, 1937, won for his work on carotenoids, flavins and vitamins: http://www.nobel.se/chemistry/laureates/1937/press.html. His biography is at (http://www.nobel.se/chemistry/laureates/1937/karrer-bio.html) Richard Kuhn, Chemistry, 1938, won for additional work on carotenoids and vitamins: http://www.nobel.se/chemistry/laureates/1937/press.html. His biography is at: (http://www.nobel.se/chemistry/laureates/1938/kuhn-bio.html) Severo Ochoa, Physiology or Medicine, 1959, won for his work on enzymatic processes in biological oxidation and synthesis and the transfer of energy. His biography is at (http://www.nobel.se/medicine/laureates/1959/ochoa-bio.html)
Melvin Calvin at Lawrence Berkeley Laboratory Melvin Calvin, Chemistry, 1961, won for his work on carbon dioxide assimilation in photosynthesis, the carbon cycle, also named "The Calvin Cycle" after him: http://www.nobel.se/chemistry/laureates/1961/press.html. It is important to mention that Andrew Benson contributed heavily to this work, and the authors of this article prefer to call the cycle, "Calvin-Benson" cycle. Calvin's biography is at: (http://www.nobel.se/chemistry/laureates/1961/calvin-bio.html). Also, an obituary may be found at: http://www.lbl.gov/Science-Articles/Archive/Melvin-Calvin-obit.html; and another one at: http://www.life.uiuc.edu/govindjee/history/obit/ObitMelvinCalvin.pdf. Jeffery Kahn writes about the establishment of the Calvin Photosynthesis Group project at UC Berkeley's Bancroft Library (http://www.lbl.gov/Science-Articles/Archive/Calvin-history-project.html). More information from the Library is available at http://bancroft.berkeley.edu/Exhibits/Biotech/calvin.html. Robert Burns Woodward, Chemistry, 1965, won for the total synthesis of chlorophyll, vitamin B12 and other natural products: http://www.nobel.se/chemistry/laureates/1965/press.html. His biography is at: (http://www.nobel.se/chemistry/laureates/1965/woodward-bio.html) George Porter, Chemistry, 1967, won for his development of flash photolysis (along with Ronald Norrish). Lord George Porter later did work on aromatic molecules and chlorophyll, energy transfer in photosynthesis and primary photochemistry of photosynthesis in femtosecond-picosecond time scale: http://www.nobel.se/chemistry/laureates/1967/press.html. His biography is at: (http://www.nobel.se/chemistry/laureates/1967/porter-bio.html) Peter D. Mitchell, Chemistry, 1978, won for his work on biological energy transfer through the formulation of the chemiosmotic theory: http://www.nobel.se/chemistry/laureates/1978/press.html. His biography is at: (http://www.nobel.se/chemistry/laureates/1978/mitchell-bio.html) Aaron Klug, Chemistry, 1982, won for development of crystallographic electron microscopy and his structural elucidation of biologically important nucleic acid-protein complexes: http://www.nobel.se/chemistry/laureates/1982/press.html. His autobiography is at: (http://www.nobel.se/chemistry/laureates/1982/klug-autobio.html) Jean-Marie Lehn, Chemistry, 1987, won for his work on mimicking natural processes such as photosynthesis and for doing the groundwork for small synthetic structures called "molecular devices": http://www.nobel.se/chemistry/laureates/1987/press.html. His autobiography is at: (http://www.nobel.se/chemistry/laureates/1987/lehn-autobio.html) Johann Deisenhofer, Robert Huber and Hartmut Michel, Chemistry, 1988, won, for determining the three-dimensional structure of bacterial reaction center using X-ray crystallography. A description of their work can be found at: http://www.nobel.se/chemistry/laureates/1988/press.html. Deisenhofer's autobiography is at: (http://www.nobel.se/chemistry/laureates/1988/deisenhofer-autobio.html). Huber's is at: (http://www.nobel.se/chemistry/laureates/1988/huber-autobio.html). And Michel's is at: (http://www.nobel.se/chemistry/laureates/1988/michel-autobio.html) Rudolph Marcus, Chemistry, 1992, won for his contributions to the theory of electron transfer reactions in chemical systems, including photosynthesis: http://www.nobel.se/chemistry/laureates/1992/press.html. His autobiography is at: (http://www.nobel.se/chemistry/laureates/1992/marcus-autobio.html) Michael Smith, Chemistry, 1993, won for his fundamental contributions to the establishment of oligonucleotide-based, site-directed mutagenesis and its development for protein studies which has become a common technique for studying photosynthesis organisms: http://www.nobel.se/chemistry/laureates/1993/press.html. His autobiography may be found at: http://www.nobel.se/chemistry/laureates/1993/smith-autobio.html Paul D. Boyer and John E. Walker, Chemistry, 1997, won for their elucidation of the enzymatic mechanism underlying the synthesis of adenosine triphosphate (ATP): http://www.nobel.se/chemistry/laureates/1997/press.html. Boyer's autobiography is at: (http://www.nobel.se/chemistry/laureates/1997/boyer-autobio.html), and Walker's is at: (http://www.nobel.se/chemistry/laureates/1997/walker-autobio.html) Ahmed H. Zewail, Chemistry, 1999, won for his studies of the transition states of chemical reactions using femtosecond spectroscopy: http://www.nobel.se/chemistry/laureates/1999/press.html. His autobiography may be found at: http://www.nobel.se/chemistry/laureates/1999/zewail-autobio.html The Light ReactionsFrom the Botany Online site at the University of Hamburg (http://www.biologie.uni-hamburg.de/b-online/e24/24c.htm), we get a brief history of the elucidation of the dark and light reactions. This is followed by an excellent description of phosphorylation and the two photosystem.The site is profusely illustrated. "Photosynthesis and Time" by Govindjee and Matej Lexa (http://www.life.uiuc.edu/govindjee/ptime/): The time sequence of the light reactions of photosynthesis are taught through a "click-on" window (with the four protein complexes in it), and through a movie. A clock covers the reactions from femtoseconds to milliseconds (recommended for all undergraduates and graduate students).
The Z-Scheme is the crux of the light reactions of photosynthesis. The scheme and its description are presented at http://www.life.uiuc.edu/govindjee/textzsch.htm. [For the scheme itself, see http://www.life.uiuc.edu/govindjee/ZSchemeG.html] June Steinberg from National-Louis University has created animations to explain the light reactions using cyclic (http://faculty.nl.edu/jste/photosynthesis.htm) and non-cyclic (http://faculty.nl.edu/jste/noncyclic_photophosphorylation.htm) photophosphorylation Light-Independent Reactions (Dark Reactions)/Calvin-Benson Cycle/Carbon CycleFrom the Botany Online site at the University of Hamburg (http://www.biologie.uni-hamburg.de/b-online/e24/24a.htm): This is a great site as the equations and the chemical structures are available. You may need to install some plug ins. From June B. Steinberg of National-Louis University (http://faculty.nl.edu/jste/calvin_cycle.htm): A beautifully done site: the steps are clearly shown. It requires downloading plugins. From Smith College, here are interesting animations showing the various steps (http://www.science.smith.edu/departments/Biology/Bio231/calvin.html) Joyce J. Diwan at Rensselaer Polytechnic Institute has a site with a very detailed explanation with numerous illustrations and links to Powerpoint presentations and test questions at http://www.rpi.edu/dept/bcbp/molbiochem/MBWeb/mb2/part1/dark.htm. Wikipedia has a good article on the Calvin-Benson cycle http://en.wikipedia.org/wiki/Calvin_cycle. The Photosynthetic Unit/Reaction Centers
The Photosynthetic Unit
from the Theoretical and Computational Biophysics Group at UIUC. "Quantum Biology of the PSU" from the Theoretical and Computational Biophysics Group at UIUC (http://www.ks.uiuc.edu/Research/psu/psu.html): It is a great site showing beautiful structures of antenna of bacterial systems. "Schematic Diagram of a Photosynthetic Unit Showing Exciton Transfer" is an animated web page from the University of Hamburg (http://www.biologie.uni-hamburg.de/b-online/library/bio201/psunit.html): It is lovely to watch it. A press release for the 1988 Nobel Prize in Chemistry" from the Royal Swedish Academy of Sciences, announcing the determination of the 3D structure of a photosynthetic reaction center (http://nobelprize.org/nobel_prizes/chemistry/laureates/1988/press.html): The text is great, but the figures are only in black and white. The Hangarter lab at Indiana University (http://www.bio.indiana.edu/~hangarterlab/courses/b373/lecturenotes/photosyn/et.html) contains a brief overview and some more complex details of the PSU. "Photosynthetic reaction centres of purple bacteria" from Scripps (http://metallo.scripps.edu/PROMISE/PRCPB.html) gives detailed structures from the PROMISE data base. "Systems Biology of Photosynthesis in Dynamic Light Environment" (http://www.e-photosynthesis.org) is a project to "offer an open web platform for modeling and reverse engineering of photosynthetic dynamism." It is a major project and includes online experiments to view results. Light-Harvesting / Antennas"Photosynthetic Antennas and Reaction Centers: Current Understanding and Prospects for Improvement," by Robert E. Blankenship at Washington University St. Louis (written while he was at ASU) (http://photoscience.la.asu.edu/photosyn/education/antenna.html). It is a well-done text with basic diagrams. We recommend it to all students. "Light Harvesting Complex II of photosynthetic bacteria" from the Theoretical and Computational Biophysics Group at UIUC (http://www.ks.uiuc.edu/Research/bio_ener/LH_2/) and also "Inter-Complex Excitation Transfer in photosynthetic bacteria " (http://www.ks.uiuc.edu/Research/psu/psu_inter.html). Excellent colored versions of the antenna complexes in photosynthetic bacteria; links are provided to PDF files of three of their research papers. "Photosynthetic Light Harvesting" by Graham R. Fleming (http://www.cchem.berkeley.edu/grfgrp/research/lightharvesting/index.htm) is a detailed slide show that is useful for visualizing much of the light harvesting process. "Organization of energy transfer networks in photosynthesis" (http://www.ks.uiuc.edu/Research/psres/) also from the TCBG (above) compares light harvesting mechanisms of anoxygenic and oxygenic photosynthetic bacteria. "Light Harvesting Complex II of Purple Bacteria" from Scripps (http://metallo.scripps.edu/promise/LH2PB.html). This site contains an excellent description and an extensive bibliography of the light harvesting complex II of purple bacteria. "Light-Harvesting in Bacterial Photosynthesis" from Glasgow University Protein Crystallography group (http://www.chem.gla.ac.uk/protein/LH2/lh2.html): Though rather brief and somewhat jarring in its color choices for the text, it does have a brief description and links to nice pictures on structure and function of the antenna system in bacteria. "Chloroplast Light-harvesting Complex II" from John Allen's site (http://jfa.bio.qmul.ac.uk/~john/webstar/Research/lhcii_chime.html): A beautiful site to visit; it is highly recommended. You can download chime and RasMol and watch the molecule while listening to Bach's Fugue in G major. Electron Transfer"Electron Transport and Energy Transduction" by John Whitmarsh at UIUC and the USDA Agricultural Research Service (http://www.ars.usda.gov/Services/docs.htm?docid=3527&page=2): It is a good review chapter from the book "Photosynthesis: A Comprehensive Treatise" edited by A.S. Raghavendra. The Z-Scheme is the description of the electron transfer in oxygenic photosynthesis. The scheme and its description are presented at http://www.life.uiuc.edu/govindjee/textzsch.htm.[For the scheme itself, see http://www.life.uiuc.edu/govindjee/ZSchemeG.html] "Introduction to Electron Transfer" by R. M. Williams of the Universiteit van Amsterdam (http://home.uva.nl/r.m.williams/Introduction%20to%20ET-30.htm): It is an excellent basic physico-chemical description of generalized electron transfer, and includes a very good exposure to the Marcus theory and the equations. "Electron Transfer in Hierarchical Photochemical Systems" from Argonne National Laboratory (http://chemistry.anl.gov/photosynthesis/hierarchical_systems_Part2.html): It deals with bacterial as well as photosystem I reactions; use of EPR, among other methods.
Electron transfer in the Photosynthetic Membrane from Robert Carpentier's web site. Robert Carpentier's site at the Université du Québec à Trois-Rivières, http://www.uqtr.ca/labcarpentier/eng/home_frames.htm, uses frames that may load very slowly, but it contains many excellent figures and is available in French and English. The section on "Photos" contains many glimpses of photosynthesizers at various conferences. Photosystems I and IICarl Bauer's lab at Indiana University (http://www.bio.indiana.edu/~bauerlab/): The site contains a nice descriptions and figures for regulation of photosystem gene regulation by oxygen and light. It also has informaiton on the origin and evolution of bacterial photosynthesis.
"Crystal structure of Photosystem I at 2.5 Å resolution" from the Institute for Crystallography at the Free University Berlin (http://userpage.chemie.fu-berlin.de/saenger/projects/ps1.html) and also, from the same site, "First x-ray crystallographic model of water oxidising photosystem II" (http://userpage.chemie.fu-berlin.de/saenger/projects/ps2.html). Highly recommended. "Plastocyanin and Photosystem I. Ru-Modification" by Örjan Hansson of Göteborg University, Sweden (http://www.bcbp.gu.se/~orjan/res/pc-ps1-e.html): It deals mainly with plastocyanin; it has references of the authors. "Shedding New Light on the Earth's Powerstation" from NASA describes the crystallizing of PS I on the space shuttle (http://www.science.nasa.gov/newhome/headlines/msad27jul98_1.htm ) and (http://science.nasa.gov/newhome/headlines/msad10may99_1.htm): It is a technical achievement. A second group of PS I crystallization experiments was destroyed in the re-entry crash of the space shuttle Columbia (http://www.asu.edu/news/faculty_students/shuttle_020703.htm) Both "Photosystem II" (http://life.uiuc.edu/~a-crofts/psiiwork.html). and "Structure and Function in Photosystem II" (http://life.uiuc.edu/~a-crofts/psiistrc.html) from Antony Crofts are good sites to learn about Photosystem II. There is some overlap in the two sites. "Photosystems I + II" from Jim Barber at Imperial College (http://www.bio.ic.ac.uk/research/barber/index.htm) shows work being done and the two photosystems in the Barber lab.
The oxygen-evolving complex from "Science Beat," a publication of the Berkeley Lab "Spinach, Or The Search For The Secret Of Life As We Know It" (http://www.lbl.gov/Science-Articles/Archive/sb/July-2004/2_spinach.html) is a nicely illustrated article in "Science Beat" from the Berkeley Lab that discusses the evolution of the oxygen-evolving complex.
Structure of Photosystem I determined using X-ray crystallography in Petra Fromme's lab at ASU "Catching some rays: Harnessing the power of photsynthesis" (http://researchmag.asu.edu/2007/03/catching_some_rays.html) discusses the evolution of Photosystems I and II. X-ray crystallography is used by Prof. Petra Fromme to determine the structure of PS I. Cytochromes and Cytochrome Oxidase"Cytochromes" from Scripps (http://metallo.scripps.edu/promise/CYTOCHROMES.html): The site contains chemical structures and descriptions of most of the cytochromes. This site is temporarily hosted by Scripps and may move in the future. "Models for Cytochrome c Biogenesis" from Washington University at St. Louis (http://www.biology.wustl.edu/faculty/models.html): It discusses three different systems for the biogenesis of cytochromes c. "Cytochrome c Oxidase" from the Theoretical and Computational Biophysics Group at the University of Illinois at Urbana (http://www.ks.uiuc.edu/Research/bio_ener/cco/): There is a basic description, and there are some key references. "Cytochrome c Oxidase" from Scripps (http://metallo.scripps.edu/promise/COX.html): It contains chemical structures; a good description; and bibliography. "The bc1-Complex Site" from Antony Crofts at the University of Illinois at Urbana (http://www.life.uiuc.edu/crofts/bc-complex_site/index.html) and Ed Berry's Cyt bc page (http://sb20.lbl.gov/cytbc1/): These sites contain the structure obtained by Ed Berry's group; Crofts page has links to coordinate data files of structures including that from J. Deisenhofer's group. "Structure-Function Studies of the Cytochrome b6f Complex" from William Cramer at Purdue University (http://www.bio.purdue.edu/people/faculty/cramer/Cramer/html/cytbf.html): Here, you will find an excellent description of this complex. ATP Synthase
Arrangement of subunits in mitochondrial ATP synthase, from John Walker's web site "ATP Synthase" by John Walker, the work that won him the Nobel Prize (http://www.mrc-dunn.cam.ac.uk/research/atpase.html): It is a simple and a beautiful page with nice colored illustrations and an incredible animations of the ATP synthase (http://www.mrc-dunn.cam.ac.uk/research/atp_synthase/rotarymech.php) and breakdown of the subunits (http://www.mrc-dunn.cam.ac.uk/research/atp_synthase/subunit.php) "Animation Movies of ATP Synthase" by Hongyun Wang and George Oster of University of California, Berkeley (http://nature.berkeley.edu/~hongwang/Project/ATP_synthase/) are great fun. Enjoy the site and learn from it. Plugins are required to view the animations. "ATP Synthase", from Antony Crofts at the University of Illinois at Urbana, includes description and crystal structure (http://www.life.uiuc.edu/crofts/bioph354/lect10.html ) "The Photosynthetic ATP Synthase: Assembly of Hybrid Complexes from Bacterial and Plant Subunits Defines Their Roles in Catalysis" by Zippora Gromet-Elhanan at the Weizmann Institute of Science (http://www.weizmann.ac.il/Biological_Chemistry/scientist/Elhanan/elhanan.html): It contains results from her research. "ATP Synthase Sites for Biochemistry" by Sandra L. Jewett, contains many good links (http://www.csun.edu/~hcchm001/wwwatp2.htm). It is a site that contains links to several sites that contain information on Paul Boyer's binding change hypothesis; movies of the ATP Synthase. It is highly recommended by the authors. C-3, C-4 and CAM (Crasulacean Acid Metabolism) Pathways"C3, C4 and CAM, Regulation of the Activity of Photosynthesis" from Botany Online at the University of Hamburg (http://www.biologie.uni-hamburg.de/b-online/e24/24b.htm) includes useful information. "How Plants Cope with the Desert Climate," by Mark A. Dimmitt, Arizona-Sonoran Desert Museum (http://www.desertmuseum.org/programs/succulents_adaptation.html): It is a basic general description of the CAM pathway written for the public. This wonderful site moves around frequently, so a search on its title might be required in the future. Brad Fiero of Pima Community College presents brief, easily understood definitions of C3, C4 and CAM photosynthesis (http://wc.pima.edu/~bfiero/tucsonecology/plants/plants_photosynthesis.htm), highly recommended for students. The Princeton/Rutgers Environmental Science Institute has posted "Global Warming Influences on C3 and C4 Photosynthesis" which is a workshop on the subject and contains information of possible interest to teachers: (http://www.woodrow.org/teachers/esi/1999/princeton/projects/c3_c4/) Chlorophyll Fluorescence"Fundamentals of Chlorophyll Fluorescence" from Optisci (http://www.optisci.com/cf.htm) is a good introduction to chlorophyll fluorescence from a commercial company. We wish more companies would post instructive information such as this, besides merely advertising their products. "Using Chlorophyll Fluorescence to Study Photosynthesis" from the Swiss Federal Institute of Technology, Zurich (ETH) (http://www.ab.ipw.agrl.ethz.ch/~yfracheb/flex.htm): It is a fairly good site with many figures and references for students. "Fluorescence as a Probe of Biological Structure and Function" from Govindjee at UIUC (http://www.life.uiuc.edu/govindjee/biochem494/biochem494a.html): You will find a bit of history; a basic description of Franck-Condon principle; basics of excitation energy transfer; some references; and a list of books. Date of site is Spring, 1999. Govindjee has also set up a page for the Kautsky Effect with a QuickTime movie showing the effect (http://www.life.uiuc.edu/govindjee/movkautsky.html).[A 3-D presentation is available at http://www.greentech.cz/lapi/about/kautsky/index.html).] "Chlorophyll fluorescence -- a practical guide" (http://jxb.oxfordjournals.org/cgi/content/full/51/345/659) is a well written review article that is available to persons that belong to schools that subscribe to the online version of the Journal of Experimental Botany. Those that cannot connect, can download the free PDF version (http://jxb.oxfordjournals.org/cgi/reprint/51/345/659.pdf). On the Use of Herbicides"An introduction to herbicides" (http://ipmworld.umn.edu/chapters/whitacreherb.htm): This site contains structures of many herbicides including Atrazine; Paraquat; Glyphosate, among others. There are also references. "Herbicide Mode-Of-Action Summary" by M.A. Ross and D.J. Childs of Crop Extension Service of Purdue University (http://www.ces.purdue.edu/extmedia/WS/WS-23-W.html) discusses the overall manner in which a herbicide affects a plant at the tissue or cellular level. "Photosynthesis Inhibitors" from the Sugarbeet Research & Education Board (http://www.sbreb.org/brochures/herbicide/photo.htm) discusses agricultural use of herbicides. The Chloroplast"Virtual Cell" is a journey into the workings of the chloroplast, from the Department of Plant Biology at UIUC (http://www.life.uiuc.edu/plantbio/cell/): This award-winning site by Matej Lexa is indeed a fun site. You can cut, zoom, turn, and really look at the "innards" of the chloroplast. Wikipedia has a good article about chloroplasts (http://en.wikipedia.org/wiki/Chloroplast). Claude W. dePamphilis at Penn State University maintains the Chloroplast Genome Database (http://chloroplast.cbio.psu.edu/) with links to information of interest to geneticists. "Biological antennae also need to be tuned..." (http://www.rhul.ac.uk/biological-sciences/AcademicStaff/Lopez-Juez/lopezlab.html) from Enrique Lopez-Juez at the University of London discusses how light and plastid signals regulate leaf and chloroplast development. There are many good photographs of Arabadopsis plants and chloroplast fluorescence and autofluorescence. Biology 4Kids (http://www.biology4kids.com/files/cell_chloroplast.html) has easy to understand explanations and figures suitable for younger students. FT Exploring is another site suitable for younger students with good explanations (http://www.ftexploring.com/photosyn/chloroplast.html). It gives simple overviews and then much more detailed information for older students. Pigments/Carotenoids"Photosynthetic Pigments" from the University of California, Berkeley (http://www.ucmp.berkeley.edu/glossary/gloss3/pigments.html): It is a nice site discussing the three major pigments of plants, algae and cyanobacteria: chlorophylls; phycobilins; and carotenoids. The site also describes the characteristics of plants and cyanobacteria (there are nice pictures of cyanobacteria, and suspensions of some algae). The International Carotenoid Society (http://www.carotenoidsociety.org/) links to the chemical structures of many common carotenoids and artcles, as well as its own newsletters. The Carotenoids Page (http://dcb-carot.unibe.ch/carotint.htm) is a beautiful site presented by George Britton. It includes several pages on: Introduction, Occurrence, Structure, and Biosynthesis of Carotenoids, followed by a list of references. The Wikipedia covers several aspects of carotenoids (http://en.wikipedia.org/wiki/Carotenoid). "Nomenclature of Caotenoids" (http://www.chem.qmul.ac.uk/iupac/carot/) are the very detailed IUPAC scientific rules for naming carotenoids. Although primarily useful only to specialists, it does gave the public a taste for the complexities of naming chemical compounds. "Photosynthetic Pigments" (http://ghs.gresham.k12.or.us/science/ps/sci/soph/energy/photosyn/pigments.htm) is a very simple explanation for K-12 students. What if chlorophyll was not the primary pigment in plants? Two sites speculate on what plant life elsewhere might be like: ‘‘Some Earth-like worlds may have foliage of colors other than green’’ from a CalTech press release (http://mr.caltech.edu/media/Press_Releases/PR12971.html) and a related illustrated article from NASA, ‘‘NASA predicts non-green plants on other planets’’ (http://www.nasa.gov/centers/goddard/news/topstory/2007/spectrum_plants.html). Rubisco (Ribulose Bisphosphate Carboxylase Oxygenase)"Rubisco: A First Look at the Mechanism" from the School of Crystallography at Birbeck, University of London (http://www.scicom.demon.co.uk/Rubisco%20proj/Title_Page.html): It is a very good site that tells you what RUBISCO is; its mechanism of action; and provides many useful references.
Structure of Rubisco showing
its four-fold symmetry. From the University of Hamburg site. "Ribulose-1,5-bisphosphate Carboxylase" from University of Hamburg (http://www.biologie.uni-hamburg.de/lehre/bza/1rxo/e1rxoe.htm): The page opens with nice colored pictures of the enzyme. The site discusses in depth the mechanism of action as well as the structure of the enzyme. It requires downloading plug-ins to view the animations. Wikipedia has a very good article on Rubisco (http://en.wikipedia.org/wiki/RuBisCO). The discovery of Rubisco is discussed in Vivienne Baillie Gerritsen's interesting article in Protein Spotlight, "The Plant Kingdom's Sloth" (http://www.expasy.org/spotlight/back_issues/sptlt038.shtml). If you like humorous top ten lists check out the "I Love Rubisco" site at (http://www.sabregirl.freeservers.com/rubisco.html). Our favorite is “#8: Space filling models look kind of like Hortas — (Star Trek (We're almost as geeky as engineers)).” Whole Plant Photosynthesis and Plant StressFree-Air CO2 Enrichment Project (FACE) cooperative projects are being set up around the globe to study the impact of carbon dioxide on plants and crops growing in the open environment. A few sites of interest are: The Face Program (http://www.bnl.gov/face/faceProgram.asp); Australian Savanna FACE (http://www.cse.csiro.au/research/ras/ozface/index.htm); EuroFACE (http://www.unitus.it/euroface/); Oakridge: A FACE experiment in a deciduous forest (http://www.esd.ornl.gov/facilities/ORNL-FACE/). There are many more; the full list is at: http://cdiac.ornl.gov/programs/FACE/whereisface.html. SoyFACE (Soybean Free Air Concentration Enrichment) is an innovative facility in Urbana, Illinois, for growing crops under higher levels of carbon dioxide and ozone: http://www.soyface.uiuc.edu. "Field Photosynthesis Measurement Systems" from New Mexico State University (http://weather.nmsu.edu/teaching_Material/soil698/Student_Material/Photosynthesis/): This site describes LICOR Gas exchange systems and their use for measuring whole plant photosynthesis. "Plant Stress" from Plantstress.Com (http://www.plantstress.com/): The site contains news from various sources that are related to plant stress. "FIFE Canopy Photosynthesis Rates Data Set Guide Document" is a technical report from Oak Ridge National Laboratory (http://www-eosdis.ornl.gov/FIFE/Datasets/Vegetation/Canopy_Photosynthesis_Rates.html). This site presents details of data on "Canopy Photosynthesis", collected in 1987 at several sites. Bacterial (Oxygenic and Anoxygenic) Photosynthesis"Introduction to the Cyanobacteria" from the University of California, Berkeley (http://www.ucmp.berkeley.edu/bacteria/cyanointro.html): It shows two nice photographs of cyanobacteria and gives a short description. Follow the in-text links and you will be rewarded with a great deal of information on the history of cyanobacteria going back more than a billion years. It also includes information on pigments, how the bacteria changed the very atmosphere we breath, and led to the existence of plants. "Purple Non-Sulfur Photosynthetic Bacteria" that supports the Bacteriology 102 course from the University of Wisconsin (http://www.splammo.net/bact102/102pnsb.html): It is a good teaching site at undergraduate level. It includes nice photos of bacterial cultures; description of bacteria and the media they are grown in. "Energy Conversion by Photosynthetic Organisms" from the Food and Agriculture Organization of the United Nations (http://www.fao.org/docrep/w7241e/w7241e06.htm): The site provides some basic information on solar energy; plant and bacterial photosynthesis; and hydrogen production by photosynthetic organisms. There are references as well. "Bacterial Photosynthesis" (http://www.biologie.uni-hamburg.de/lehre/bza/photo/ebacphot.htm) from the University of Hamburg is very detailed and requires plug-ins. Evolution of Photosynthesis"Origin of Microbial Life and Photosynthesis " from Carl Bauer at Indiana University (http://www.bio.indiana.edu/~bauerlab/origin.html): The site has a clear text, beautiful evolutionary trees, and the photosynthetic gene cluster of heliobacteria. "Evolution: When Did Photosynthesis Emerge on Earth?" by David J. Des Marais at NASA Ames Research Center (http://edmall.gsfc.nasa.gov/aacps/news/Photosynthesis.html) and his article "Evolution of Earth's Early Biosphere and Atmosphere" (http://astrobiology.arc.nasa.gov/palebluedot/abstracts/ddm.html): The first text consists of extracts from a paper by Jin Xiong and coworkers, published in Science (Sep. 8, 2000). "The Manganese-calcium oxide cluster of Photosystem II and its assimilation by the Cyanobacteria" by James D. Johnson (http://www.chm.bris.ac.uk/motm/oec/motm.htm) discusses the evolution of the oxygen-evolving complex in bacteria. It is very complete and contains many good illustrations and chemical structures. "Geobiologists Solve "Catch-22 Problem" Concerning the Rise of Atmospheric Oxygen" (http://pr.caltech.edu/media/Press_Releases/PR12927.html) is a news release from Caltech that discusses some puzzles about the early evolution of cyanobacteria and their ability to produce oxygen without poisoning themselves. Now for something completely different: "An animated depiction of the origins and diversification of oxygenic photosynthesis" done in claymation by Felisa Wolfe-Simon and friends (http://www.ironlisa.com/claymation.html). Photosynthesis and the Environment / Global Climate Change & Warming"Greenhouse Gases and Society" by Nick Hopwood and Jordan Cohen from the University of Michigan (http://www.umich.edu/~gs265/society/greenhouse.htm) covers many aspects including the Greenhouse Effect, greenhouse gases, effects on the environment and society. Many good figures. "The Decade After Tomorrow: Modeling Global Climate Change at Berkeley" (http://sciencereview.berkeley.edu/articles.php?issue=7&article=greenhouse) by Kristen DeAnglis discusses work being done at UC-Berkeley. It is a good general article that discusses the problem and its complexities. "Global Climate Change Policy Book" (http://www.whitehouse.gov/news/releases/2002/02/climatechange.html) from the Bush White House describes his administrations plans for dealing with global climate change. Curious about the Kyoto Protocol? Here is the full text (http://unfccc.int/resource/docs/convkp/kpeng.html). "Tropical Forests and the Greenhouse Effect: A Management Response" by Norman Myers and Thomas J. Goreau (http://www.ciesin.org/docs/002-163/002-163.html): It is a 1991 paper; has text and references, but no figures "Photoinhibition in Antarctic Phytoplankton by Ultraviolet-B Radiation in Relation to Column Ozone Values" from NSF's Office of Polar Programs (http://www.nsf.gov/od/opp/antarct/ajus/nsf9828/9828html/j1.htm): It is a short article, has some references, and deals with ozone-related problems. "Helping Ocean Algae Could Beat Greenhouse Effect" by Bill Rosato describes how adding iron compounds to the water may help algae reproduce faster, thus removing more CO2 from the atmosphere, (http://www.climateark.org/articles/2000/4th/heocalgc.htm). It is a 2-page news item. "The Colors of Life" from NASA demonstates how satellites are tracking photosynthesis in the world's oceans to determine how well phytoplankton utilize carbon, http://www.gsfc.nasa.gov/gsfc/earth/pictures/2001/0327colors_of_life/carbon.htm. "Photosynthesis, Trees, and the Greenhouse Effect" is a lesson plan for teachers from the National Geographic (http://www.nationalgeographic.com/xpeditions/lessons/08/g68/brainpopphoto.html). Rhodopseudomonas palustris (http://genome.jgi-psf.org/finished_microbes/rhopa/rhopa.home.html) is a bacterium with some interesting abilities that may be useful for our environmental future: it can sequester carbon dioxide and can produce hydrogen. A news release from the Ohio State University also discusses this useful bacterium (http://researchnews.osu.edu/archive/micromet.htm). The EPA (Environmental Protection Agency) has a kids site on the "Greenhouse Effect" (http://www.epa.gov/climatechange/kids/greenhouse.html) with simple explanations and links. "Mimicking Nature's Engine" by Erik Ellis discusses some of the work of the ASU Center for the Study of Early Events in Photosynthesis (http://researchmag.asu.edu/stories/mimick.html). It is a news item with a short basic description. The Swedish Consortium for Artificial Photosynthesis (http://www.fotomol.uu.se/Forskning/Biomimetics/consortium/index.shtm) discusses their research goals of using artificial photosynthesis to produce hydrogen. "Artificial Photosynthesis" by the Wasielewski group at Northwestern University (http://www.chem.northwestern.edu/~wasielew/res/ap.htm) discusses their work designing an artificial reaction center. "Energy at the Speed of Light" by Andrew Gathman at Penn State University (http://www.rps.psu.edu/0009/energy.html): It is a news item worth reading. "Theory and Modeling of Biological Nanodevices" by Klaus Schulten at the UIUC (http://www.foresight.org/Conferences/MNT8/Abstracts/Schulten/ ) contains a brief summary of using photosynthesis to develop nanodevices‹an area of research that is pushing the envelope of photosynthesis research. "Reinventing the Leaf" by Philip Ball in Nature: Science Update (http://www.nature.com/nsu/991007/991007-3.html). It is a news item worth reading. Bioenergy and PhotosynthesisSteve Chu, the Nobel Prize laureate
from Lawrence Berkeley lab writes about the promise of photosynthesis in "Worldwide
Energy Crunch: "Photosynthesis and Bioenergy" (http://sols.asu.edu/grad/igels/photosynthesis.php) briefly describes new graduate programs at ASU that will tackle issues for the future. Stay tuned. Oakridge National Laboratory has several programs to study bioenergy. A couple interesting web pages are some Frequently Asked Questions (http://bioenergy.ornl.gov/faqs/index.html) and the bioenergy cycle (http://bioenergy.ornl.gov/papers/misc/bioenergy_cycle.html). The newly established Center for Bioenergy and Photosynthesis at ASU contains numerous links to this developing area of research (http://photoscience.la.asu.edu/CB&P/). The center is a large multidisciplinary group and incorporates the former “ASU Center for the Study of Early Events in Photosynthesis.” The ASU Research Magazine has two articles about recent activities. "Catching some rays: Harnessing the power of photsynthesis" (http://researchmag.asu.edu/2007/03/catching_some_rays.html) discusses the history and evolution of photosynthesis and its potential to develop sustainable energy sources. "Bacteria for biofuel" (http://researchmag.asu.edu/2007/03/bacteria_for_biofuel.html) discusses current projects to develop biofuels from cyanobacteria, a more rational approach than using food crops or rainforests. |
Author Contact Information : Larry Orr Govindjee This article is based on a Review published |
| 14 September 2007 |
