ENERGENICS   PHOTOSYNTHESIS / RESPIRATION

                                                             Pages   280 - 340
 

Metabolism - total of all chemical reactions in a living system.

Anabolism- energy poor molecules are transformed into energy rich molecules absorbing energy

Catabolism - energy rich molecules are transformed into energy poor molecules releasing energy

Autotrophic/Heterotrophic  type of nutrition. Look up definition in glossary of text.

ATP are molecules which are able to transfer energy directly to reactant molecules during chemical reaction within a cell.

 ATP -    chemical compound adenosine  tri?phosphate.

             The adenosine  part of ATP is the main body of   the molecule; it is
             composed of adenine (also a component of  DNA) and a simple sugar, ribose.

             The tri- part of the name refers to the fact that ATP has three phosphate
             groups linked to the main adenosine body of the molecule.

Cyclic phosphorylation:                                 work

                                    ATP ---------------------------------> ADP + (P) + E
 

                                   ADP + (P) + E  ------------------------ > ATP

                                                        respiration

                               C6H12O6  +  ATP     ---------->     C6H11O6PO4     +   ADP
 
 

  Refer to figures handed out in lecture.
 
 
 
 
 

 ATP is not the only molecule that may be used to transfer biological energy from  energy rich molecules. Three other molecules (coenzymes) are nicotinamide adenine  dinucleotide (NAD+) and flavin adenine dinucleotide (FAD). During photosynthesis the coenzyme NADPH2 is used in place pf NADH2

See Handouts From Class
 

Both of these molecules may be reduced by various chemical reactions  (respiratory).

 Example:        NAD + CH3CH2OH  -----> NADH+H + CH3COOH

 Note the NAD molecule removes  hydrogen atoms.

Example:         NADH+H  +  CH3COCOOH ---> CH3CHOHCOOH + NAD

Note the NADH+H  molecule donates  hydrogen atoms.

These are called coenzymes: coenzymes you need to know are NAD, FAD and NADP
 

These two reduced forms (NADH+H and FADH2) can also  be transferred to the cytochrome system in a mitochondria where they will be oxidized with the energy  being released converted into working ATP.   Oxygen is required for this process.
 

                           O2    +    ADP + P + NADH  -----> NAD + ATP +   HOH

   ATP can be synthesized from energy rich molecules in two ways.  (Look above)

           1.Energy is directly taken from a substrate to form ATP from ADP and (P).
                This is called substrate phosphorylation.  (single step process)
           2.Energy is taken from an energy rich molecule by reducing  NAD+ or FAD
              to NADH+H or FADH2 These molecules move to a cytochrome system in a
              mitochrondrion where the reduced forms NADH+H or FADH2 are oxidized
              to NAD+ or FAD and the accompanying energy being transferred to make
              ATP. This is called oxidative phosphorylation.  (two step process)

During photosynthesis the coenzyme NADPH + H is used in place of NAD + H
 

See Handouts From Class
 

        Cellular respiration is the chemical process (oxidation) of breaking down energy
        rich molecules (C6H12O6) into energy poor molecules (CO2 + H2O) with the
        energy being converted into biologically usable energy - ATP. This is a slow
        process and requires several reactions which can be categorized into steps.

    Aerobic respiration has three stages: glycolysis, citric acid cycle (Krebs
    cycle), and electron transport system. A limited amount of ATP is produced
    directly from specific steps in gylcolysis and the Kreb cycle (substrate
    phosphorylation). However, the bulk of  ATP will be formed by the formation of
    NADH+H and FADH  via glycolysis and the Kreb cycle which then must move to
    the electron transport system to be transformed into ATP using oxygen (oxidative
    phosphorylation). The overall equation for aerobic respiration is:
 

                                      36ADP + 36P + C6H12O6 + 6O2 ------> 6CO2  + 6H2 O + 36 ATP
 
 

 Glycolysis is a series of chemical reactions which occur in a step by step process.
 During each step an organic molecule is  converted to a different molecule
 associated  with an energy change. These steps will resemble a "road map "showing
changes in organic molecules  and the involvement of biological energy. The two  forms
of chemical biological energy involved during these reactions are ATP and
 NADH + H.

Examine the Glycolysis and prepare an  input/output box in your notes
indicating  what enters and leaves this pathway.  Include the organic molecules and
the biological energy.

See Handouts From Class
 

                                                              input/output box
                                                    (write in your notes and learn)

  Organic Molecules:   Input: glucose   Output: 2 pyruvic acid molecules
  Energy Output: ATP ( 2 used during activation and 4 synthesized = net of 2)
  NADH (2 formed which may be converted into 2 ATP per NADH
  thus a total of four ATP may be formed by these coenzymes)

  Glycolysis produces a net of 2 ATP directly and 4 through NADH, a total of 6
      ATP per glucose.

              Substrate phosphorylation refers to the ATP made directly.
              Oxidative phosphorylation refers to the ATP synthesized through the
              NADH
 

Aerobic respiration also involves the Kreb Cycle.  During this process pyruvic acid
which was formed during glycolysis is going to be oxidized into carbon dioxide and
 water producing ATP directly (by substrate phosphorylation)  and the coenyzmes
(NADH + H and FADH) which  form ATP via cytochrome  system (oxidative  phosphorylation).

Prepare an  input/output box in your notes indicating  what enters and leaves the Kreb Cycle pathway.  Include the organic molecules and the biological energy.

See Handouts From Class
 
 

Organic Molecules :   Input: pyruvic acid   Output: 3 carbon dioxide molecules

 Energy Output ? ATP - 1 via substrate phosphorylation,   NADH - 4 formed which
 may be converted into 3 ATP per NADH, FADH – 1 formed which may be
 converted into 2 ATP per FADH.  Total of 15 per Kreb cycle.
 
 
 

The conversion of NADH, FADH into NAD,FAD forming ATP involves the cytochrome
system utilizing oxygen. Cytochrome Electron Transport System

See Handouts From Class
 

Prepare an  input/output box in your notes indicating  what enters and leaves the CETS  pathway.

No organic molecules are broken down.

NADH and  FADH  may enter the chain which provided energy to drive a proton pump. This pump forms a proton gradient in the crista lumen of the mitochondrion which allows the enzyme ATPase to to form ATP.

ADP + P  -----à     ATP  + HOH

The end receptor of the hydrogen ions is oxygen. (oxygen is consumed)

Therefore this set of reactions is called oxidative phosphorylation.
 

 Energy Output – 3 ATP per NADH + H and 2 ATP per FADH + H
 
 

The NADH+H produced during glycolysis has to move through mitochondrion; therefore ATP is required for active transport. This is why these NADH + H produces only a net of two ATP where NADH + H is mitochondrion produces three ATP.
 

Anaerobic Respiration

Includes  glycolisis; however the pyruvate is reduced into ethanol and
carbon dioxide.  The reducing power comes from the NADH + H produced during glycolysis; therefore they can not be used to produce ATP via oxidative phosphorylation.
 

                                      2ADP + 2P + C6H12O6  ---> 2CO2 + 2C2 H5OH + 2 ATP
 
 

See Handouts From Class

Examine glycolysis and the reduction of pyruvate into carbon dioxide and ethanol.
 
 

                                                              input/output box
                                                    (write in your notes and learn)

  Organic Molecules -   Input: glucose   Output: 2 ethanol and 2 carbon dioxide
  Energy Output - ATP ( 2 used during activation and 4 synthesized = net of 2)
  NADH (2 formed which are used to reduce pyruvate)
 
 

Other catabolic reactions producing energy and carbon skeletons.

Pentose Phosphate Pathway

    Forms energy and ribose sugars for DNA, RNA synthesis.
 
 

See Figures in text involving Pentose Phosphate Pathway
 
 

Respiration of Lipids
 

Lipids (triglycerides ae hydrolyzed into glycerol and fatty acids.
       Glycerol can be converted into pyruvate and metabolized.
       Fatty Acids can be broken down into acetates (2 carbon molecules) via beta
       oxidation and enter the Kreb Cycle.
 

See Figures in text involving break down of fat
 

Heat-generation reactions

Thermogenic reactions or cyanide resistant reactions.

If NADH breaks down to NAD, a large amount of energy is liberated; and if this powerful exergonic reaction is not coupled to any endergonic reaction, all the energy is converted to heat. (ex. Skunk cabbage)
 

See Figures in text involving thermogenic reactions
 
 

                                                      Photosynthesis

Photosynthesis is a chemical process where energy poor molecules (CO2 and
H2O) are converted to energy rich molecules (C6H12O6) with the energy to
synthesize the energy rich molecule coming from light. The overall equation is:

                                                        light
                         6CO2 + 6H2O ------------------->       C6H12O6 + 6O2
                                                    Chlorophyll

Photosynthesis  has two  stages: light dependent reactions and the light independent  reactions (dark reactions) or Calvin Cycle .  ATP and NADPH+H are produced during the light dependent  reactions and carbon dioxide is fixed into glucose during the dark reactions.

Light Dependent Reactions:

Light – red (P680, P700) and blue are utilized

Pigments:  chlorophyll “a”, chlorophyll “b”,  carotenoids (carotene and
                   xanthophylls); water insoluble

                    anthocyanin is a red water soluble pigment not used during
                    photosynthesis.

See Handouts From Class
 

Absorption Spectrum
 
 

See Handouts From Class
 
 
 

Action Spectrum
 
 
 

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Photosystem I and Photosystem II are used to convert light energy into chemical energy  (1) by passing electrons through a cytochrome system which generates ATP; (2) reducing NADP  to NADPH.
 

Photosystem I:

Uses P700 chlorophyll (absorbs red light at 700 nm range) to excite electrons
Electron acceptor  “X Fe4S4”
This reduced molecules passes electron to ferrodoxin  which reduces NADP  to NADPH

Phototsystem II:

Uses P680 chlorophyll (absorbs red light at 680 nm range) to excite electrons
Electron acceptor  “phaeophytin”
Passes electron down a cytochrome system (b6/f) generating ATP
Plastocyanin passes electron to P700 chlorophyll
Water is split to replace the electrons lost.
 

See Handouts From Class
 

Input: light,   ADP,  NADP, water

Output: ATP, NADPH+H, oxygen

Photophosphorylation ATP is generated utilizing light energy (compare to substrate and oxidative phosphorylation)

How does cytochrome system generate ATP?   Uses chemicl energy from NADH or FAD received from glycolysis
                                                                               or Kreb.
 

See Handouts From Class
 
 
 

Compare this to how ATP was generated during oxidative phosphorylation.

Two parts to oxidative phosphorylation

1) Uses energy from NADH and FADH (formed during glycolysis and Krebs) to drive proton pump
2) The proton gradient formed by pump runs the ATPase enzyme
     which generates ATP.

See Handouts From Class

Two parts to photophosphorylation

1) Uses energy from light  to drive proton pump forming a hydrogen
     gradient
2) The proton gradient formed by pump runs the ATPase enzyme
     which generates ATP.

Review aerobic respiration using input/out boxes.
The carbon, hydrogen and oxygen atoms of glucose are converted into which molecules?

Some energy is transferred into ATP directly.

Some energy is transferred into NADH and FADH which will be oxidized to generate ATP (oxidative phosphorylation).

NADH and FADH (from glycolysis and Krebs cycle) supplied the power to form ATP.

Light supplies the power to form ATP  during photophosphorylation.
 
 

Light Independent, Dark or Calvin Cycle Reactions
 

Utilize the energy (ATP and NADPH+H) formed in the light to reduce carbon dioxide into glucose
 

The main enzyme is RUBISCO     -  RuBP  carboxylase
 

See Handouts From Class
 
 

                                                                         12 ATP                                 12 ADP

Draw:            6  RuBP   +   6 CO2      -----------> 12  PGA----------> 12 PGAL
                      5 carbon           1 carbon                   2 (3carbon molecule)

                                                                          12 NADPH+H            12 NADP
 
 

See Handouts From Class
 
 
 

2 of the PGAL move through glycolysis backwards to form 1 glucose molecule and the remaining 10 PGAL recycle to reform the RuBP
 
 
 
 

input: carbon dioxide, (sugar ribulose biphosphate is recycled), ATP, NADPH+H

output: glucose, ATP, NADP

Light is not directly needed for this process; however, the ATP and NADPH+H formed during the light reactions are required.
 

RUBISCO -  RuBP   This enzyme also fixes oxygen into phosphoglycolate etc.
                       (photorespiration). Therefore, there is competition by oxygen and
                       carbon dioxide for this enzyme.   inefficient
 

Where do these reactions occur?

Chloroplast (double membrane structure with liquid part stroma and membrane sacs called grana)
 
 
 

Light dependent reactions occur in the thylakoids of grana and stroma;  the Calvin Cycle enzymes located in the stroma.

The above reactions are called C-3 because the first product is PGA

Another type of carbon fixing pathway is termed C- 4
                                 (found in  tropical grasses – corn, sedges, etc)
The pathway uses an enzyme PEP carboxylase to fix carbon dioxide into  malate

                                 PEP   + CO2   --------------->   malate  (OAA)
                            3 carbon          1 carbon                      4 carbon
 

PEP carboxylase has a high affinity for carbon dioxide and does not show photorespiration, therefore no competition with oxygen.
 
 

PEP carboxylase has a high affinity for carbon dioxide; the carbon is fixed in the mesophyll cells of the leaf;
 

See Handouts From Class
 

Plants (C-4 plants) have Kranz anatomy (well developed bundle sheath cells); the malate (OAA) is formed by PEP carboxylase in the mesophyll cells. The malate moves into the bundle sheath cells where it is decarboxylated forming carbon dioxide. In the bundle sheath cells the carbon dioxide(high amounts) is fixed into glucose via RuBP carboxylase. Basically carbon dioxide can be efficiently fixed in mesophyll cells by PEP carboxylase (C-4), move to bundle sheath cells in the form of malate, decarboxylated in bundle sheath cells forming a high concentration of carbon dioxide which is converted into glucose  by RuBP (C-3 metabolism).
 

See Handouts From Class
 
 

 Why have a C-4?

1. PEP carboxylase is much more efficient in fixing carbon dioxide

2.  C-3 plants have photorespiration where oxygen competes for the RuBP carboxylase enzyme oxidizing PGA into carbon dioxide (wasteful reaction)
(see figure)
 
 
 
 
 

See Handouts From Class
 
 

CAM plants uses the metabolism involved in C-4 plants but the timing for each pathway is
different to conserve on water (stoma are closed during day).
                        (desert succulents Agave, Cacti. Bromeliads, Orchids)
These plants will have stomata closed during day and open at night to conserve on water.  During the night (stomata open, carbon dioxide may enter) PEP carboxylase fixes carbon dioxide into malate (C-4 metabolism).  During the day (stomata closed to conserve water, carbon dioxide may not enter) malate is decarboxylated forming carbon dioxide.   RuBP  fixes the carbon dioxide into glucose via C-3 pathway.
 
 
 
 
 

See Handouts From Class

FILL OUT THE TABLE BELOW COMPARING THREE TYPES OF LIGHT INDEPENDENT REACTIONS

                                                             C-3                               C-4                                  CAM

Main
Carboxylating
Enzyme

Location
Of Enzymes

When are Stomata
Open

When are Carboxylating
Enzymes Functioning

When are Light Dependent
Reactions Functioning

Type of Plants Exhibiting
This Pathway
 
 
 

Environmental Requirements  for Respiration and Photosynthesis
 

Light Quality  -  Which wavelengths reach the plant will have an effect on how the plant grows.

Light Quantity  - The intensity of light which reaches the plant will have an effect on plant growth.

     Light compensation point – The light intensity where photosynthesis =
     respiration.

     Light saturation point – The light intensity where an increase in light intensity
     will not cause an increase in the photosynthetic rate.
 

See Handouts From Class
 

Temperature

   Between the temperatures 5 and 25 C an increase of 10 degrees causes a doubling
   of the reactions.  (Both photosynthesis and respiration)

Oxygen -  Low oxygen concentration will cause respiration to switch from aerobic to
                 anaerobic respiration.  During anaerobic respiration the pyruvic acid
                 formed during glycolysis will be reduced into ethanol and carbon dioxide.
                 The reason for this is that lack of oxygen inhibits the cytochrome system
                 which oxidized NADH to NAD; therefore, in the absence of oxygen the
                NAD will not be generated to be used to reduce the organic compounds of
                the Krebs cycle.