Chapter 5.  Photosynthesis                                                                             BI 101 Davison Fall 2006

Photosynthesis is an energy conversion converting light energy to chemical bond energy 

Organic molecules are the key building blocks and energy stores for life.  Plants make (synthesize) their own organic molecules from  inorganic atmospheric CO2 .  To do this plants trap light energy to drive the metabolic pathways we call photosynthesis. 

When eaten by animals or decay organisms, plants provide the organic molecules needed by those forms of life.  You also need plant-made organic molecules to supply the building blocks and energy needs required for growth and life itself.  You get these organic molecules by eating the plant that created them or by eating organic molecules originally found in plants but  now reside in some food product you may be about to eat.  Thus plants are the primary source and sole producer of original organic food. 

I.     Historical Perspective 

In the early 1600's it was a widely held notion that plants absorbed food from the soil (Aristotle wrote that leaves in fact were mere shading devices providing a comfort zone for roots; everyone new adding manure and humus to soil resulted in plants growing faster and growing larger.) 

1.   Mid 1600's van Helmont - a Belgian physician, devised simple experiment in which he grew a willow tree seedling for five years in a bucket containing a known quantity of soil.  The tree gained roughly 195 pounds while the soil lost less than one pound. 

                                     5 lb. willow                          200 lb. 

                                                         ----------->
 
                                                               5 yr
                                      
200 lb soil                         199.8 lb soil 

 Van Helmont concluded wrongly that plants obtain food from water.

If most of the dry material (dry biomass) didn’t come from soil or rain water, then where did it come from?  It seems unlikely based on perceptions limited to our bodily senses, but tremendous tonnage of biomass accumulates each day from thin air.  Photosynthesis is the process by which this happens—that seemingly weightless CO2 is pulled from the air and accreted into an organic form, the form of carbohydrates.  Water from the soil plays an important role in supplying atoms of Hydrogen that are required in building carbohydrates (so Van Helmont wasn't entirely incorrect).  Soil also provides inorganic nutrients that supply the other elements used in constructing other organic molecules such as amino acids and nucleotides (all but C, H, & O are from the soil, recall the mnemonic device for the chemical elements of life “C HOPKNS CaFe Mg B Mn CuZn Mo Cl Na”). 

2.  1700's Joseph Priestly (England) - put things under bell jars.  Mouse kept alive in bell jar with plant.
King of England presented Joseph Priestly with a medal and declared
"for these discoveries we are assured that no vegetable grows in vain... but cleanses and purifies our atmosphere."  Thus before 1800, it was a known fact that plants give off oxygen gas needed in the respiration of animals.  Why the oxygen was given off, or the process by which plants produce oxygen gas, remained a mystery. 




II.        The Chemistry of Photosynthesis:   

1.  Light Dependent Reactions  
2.  Light Independent Reactions

1.  Light Dependent Reactions - require light  (see section 5.2, p. 76-77)

Within the range of visible light (recall that white light consist of a spectrum of colors-energy waves-ROY G BIV) photosynthetic plant pigments capture (absorb) energy from the blue/violet and red portions of the spectrum while green and yellow light is reflected.  

The “z-scheme” (fig. 5.6a) is the model that best explains these reactions.  These reactions occur on the inner membranes of the chloroplast; these membranes hold the pigments and other molecules involved.  Upon the inner chloroplast membranes photosynthetic pigments are arranged into discrete patches called photosystems (illustrated as green globs embedded in the phospholipid bilayer shown in figs. 5.7).   It is these pigment molecules (largely chlorophyll) that absorb light energy.  Absorbed light energy, from the blue and red portions of the spectrum predominantly, is transferred to electrons causing the electrons to leave the photosystems.  These removed electrons give up their energy absorbed from the sun to produce ATP during the Electron Transport Chain (more about ETC below).   To replace electrons removed from the photosystems, electrons are taken from water molecules.  Removing electrons from two H2O’s results in the water molecules splitting into four H+ and two O atoms.   The O atoms combine forming O2 as a waste molecule or byproduct.  This process of splitting water molecules is called photolysis.

            Electron transport chain – a difficult concept, but go with the following analogy:  The electrons that begin the electron transport chain have potential energy stored from the sun just as water behind a dam stores potential energy from the sun (Recall the sun is needed to elevate water, via evaporation and rainfall, from sea level to the level of the dam.  Similarly, sunlight elevates the potential energy of electrons).  The electrons are transported between various molecules in the membranes of the chloroplast such that they give up their potential energy in a form that is used to add a phosphate to ADP, thus making ATP.  This is very much like the water behind a dam giving up its potential energy to generate electricity as it (the water) is transferred via the pull of gravity through the turbines in the dam.  A more detailed explanation is found in your text (a buildup of hydrogen ions that form from the ETC and from photolysis is used as a force that makes ATP just as water behind a dam is used as a force to make electricity;  the concentrated hydrogen ions flow across the membrane to where they are less concentrated through a transport protein that then makes ATP--hydrogen ions passing through the special transport protein that makes ATP is like water passing through a turbine that makes electricity).  

In summary of the Light Dependent Reactions, O2 has been produced as a byproduct and light energy has been converted into the chemical bond energy of ATP.  But ATP is not food, it is incapable of being stored, and thus photosynthesis is not yet complete.

2.  Light Independent Reactions- do not require light. See fig. 5.8.

              =Carbon fixation – the conversion of inorganic CO2 into carbohydrates.

            =Calvin cycle, Calvin-Benson cycle, or C3 cycle.

                         1940's - Calvin & Benson - used 14C radioactive isotope and traced the path of carbon fixation - conversion of inorganic CO2 into organic carbohydrates.  Today this reaction series bears their name as the Calvin-Benson Cycle or Calvin Cycle.

                       The actual synthesis reactions that produce carbohydrates during photosynthesis do not directly require light.  These so-called light independent reactions do however require the ATP made during the light reactions.  Without ATP, no energy would be available to drive the carbon fixation process.  Carbon and Oxygen atoms from atmospheric CO2 and the H+  derived from water pulled from the soil are combined to make carbohydrates during the Calvin Cycle.  All the chemicals involved are dissolved in the solution that fills the chloroplast, thus the internal membranes of the chloroplast are not directly involved in the Light Independent Reactions.  As carbohydrates accumulate they are transported to other plant organs or stored as starch granules inside the chloroplast.  Some of the products of photosynthesis (i.e. carbohydrates) are eventually converted to lipids or combined with nitrogen taken from soil and used to construct proteins and nucleic acids (e.g. DNA).

In summary of the Light Independent reactions:

·        consumes six molecules of CO2  to produce one molecule of glucose, C6H12O6

·        consumes ATP made during light reactions (ATP is the energy source that directly drives the carbon fixation reactions)


III.       Importance of Photosynthesis:

Fossil fuels - plants from 350 MYA (Carboniferous Period) died, were covered with sediments, and became coal.

Food - 250,000 plant species, yet remarkable few used as human food.  Only 150 crop species.  The big four are:  rice, wheat, corn, & beans  (the first 3 are grasses).

Removes CO2 - reduces greenhouse gas.  – today only 0.033% CO2 ;  earth billions of years ago had much higher CO2 levels, much of this early CO2 has been incorporated by plants into biomass (or into fossil fuels).  Our removal of forests and combustion of fossil fuels returns CO2 to the atmosphere and results in: 1) theoretically with all other factors removed, faster plant growth & 2) global warming.

Created O2 rich atmosphere (about 21 % O2)
Lead to the formation of the ozone (O3) layer which filters harmful UV radiation).  
3O2 ---> 2O3 in stratosphere

An O2 rich atmosphere and ozone layer are prerequisites for life on land---a rather recent occurrence.  Only 10% of the time that life has been on earth has there been substantial terrestrial life.  For the first 3 billion years life on earth was essentially limited to aquatic environments and only in the last 400 million years has terrestrial life appeared.  Photosynthesis performed by algae and seaweed changed remarkably the earth’s atmosphere making the life on land possible.

IV.       Chemosynthesis - some bacteria use chemical energy to fix CO2

            Not all organic molecules are created by plants and photosynthetic algae, a minor amount of primary production occurs by chemoautotrophs, bacteria which use chemical bond energy of iron and sulfur compounds to fix carbon dioxide.  Chemoautotrophs are important in nutrient cycling.

V.        Additional Vocabulary:

            Autotrophs – organisms that create their own food by fixing CO2 into an organic form: requires outside energy source – sun in the case of photoautotrophs (ex. Algae, plants) & inorganic chemical energy in the case of  chemoautotrophs (some bacteria).

            Heterotrophs – organisms that feed on food already formed by autotrophs.  (ex. some non-green plants, all animals, some bacteria, all fungi)

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