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2.

Metabolism of microbes

• Energy conservation
•Application in biosynthesis
Edited by Dr. Harbant (April, 2012)
Energy conservation
Fueling Processes in chemoorganotrophs
◆ Chemoorganotrophs differ in the types of electron acceptors they
use in their various energy-yielding processes
◆ The process involving exogenous (external) acceptors in the
electron transport chain is termed respiration
✳ Final electron acceptor is oxygen, the process is termed aerobic
respiration
✳ Final electron acceptor is not oxygen, the process is termed
anaerobic respiration
◆ The electron transport system uses the donated electron to make
ATP via oxidative phosphorylation
◆ When the electron transport system is not used and endogenous
(internal) acceptors are employed, the process is termed
fermentation and ATP is formed via substratelevel phosphorylation
Aerobic respiration

◆ Aerobic catabolism can be divided into three different stages:


✳ Stage I - larger molecules are broken down into their constituent
parts with little energy released
✳ Stage II - either an aerobic or anaerobic process whereby even
simpler molecules are produced as is ATP as well as NADH and/or
FADH2
✳ Stage III - complete oxidation of molecules under aerobic conditions
to form CO2 as well ATP, NADH, and FADH2 (the latter two
molecules generate even more ATP through the electron transport
system)
◆ Some of the pathways in metabolism are amphibolic, i.e., they
function both catabolically and anabolically
Breakdown of glucose
◆ Microbes use a number of pathways to
catabolize glucose to pyruvate, among which
include the following:
✳ Glycolysis
✳ Pentose phosphate pathway
✳ Entner-Doudoroff Pathway
Glycolysis (Embden-Meyerhof or Glycolytic
Pathway)
✳ Most common pathway found in all major groups of microbes
✳ Functions in the absence or presence of oxygen
✳ Located in the cytoplasmic matrix of both procaryotes and eucaryotes
✳ Divided into two parts
● Six carbon stage - glucose (6 carbon [C6] molecule) is phosphorylated twice
and converted to fructose 1,6-bisphosphate [C6]
– Input of 2 ATP molecules
– No energy produced
● Three carbon stage - catabolism of fructose 1,6-bisphosphate [C6] to two
molecules of pyruvate [C3] and generate per pyruvate
– One molecule of NADH
– Two molecules of ATP by substrate-level phosphorylation
✳ Substrate-level phosphorylation is the synthesis of ATP by coupling ADP
phosphorylation with an exergonic reaction
Overview of glycolysis

● Six-carbon stage
– ATP + Glucose [C6] = glucose-6-phosphate [C6]
– Glucose-6-phosphate [C6] + ATP = fructose-1,6-bisphosphate [C6]
● Three-carbon stage
– Fructose-1,6-bisphosphate [C6] cleaved into glyceraldehyde-3-
phosphate [C3]
– Glyceraldehyde-3-phosphate [C3] converted to pyruvate [C3] + 2 ATP +
NADH (this process occurs twice - balance the carbon atoms!!! 2 x C3 =C6)
✳ Summary of glycolysis
● 2 pyruvate molecules
● 2 ATP molecules - sum total: 2 used to begin glycolysis, 4 produced by
substrate-level phosphorylation
● 2 NADH molecules
PPP Glycolytic pathway
Pentose Phosphate Pathway (Hexose
monophosphate pathway)
✳ May be used simultaneously with glycolysis
and Enter-Doudoroff pathway
✳ Operates under either aerobic or anaerobic
conditions
✳ Important in both catabolism and
biosynthesis
Summary of PPP
Summary of pentose phosphate pathway
● NADPH formed serves as a source of electrons for
reduction of molecules during biosynthesis
● Four- and five-carbon sugars are synthesized that
will be used for a variety of purposes
● Intermediates are used to produce ATP
● Serves as a catabolic pathway for five-carbon
sugars
● More important as an anabolic pathway than as a
catabolic pathway to produce energy
Entner-Doudoroff Pathway
✳ Begins the same way as the pentose phosphate
pathway to produce glyceraldehyde-3-phosphate
✳ Glyceraldehyde-3-phosphate catabolized to
pyruvate via the 3-carbon stage of glycolysis
✳ Total yield per glucose molecule
● One ATP molecule
● One NADH molecule
● One NADPH molecule
TRICARBOXYLIC ACID CYCLE
◆ Tricarboxylic Acid Cycle (Krebs Cycle, TCA Cycle, Citric Acid Cycle) is widely distributed
among microbes

◆ Comprised of five basic steps:


✳ Oxidation of pyruvate [C3] to acetyl-coenzyme A (AcCoA) [C2] with the release of
CO2 and the formation of NADH

✳ AcCoA [C2] condenses with oxaloacetate (OAA) [C4] to form citric acid (citrate) [C6],
the first compound of the 6-carbon stage

✳ Through a series of reactions in the six-carbon stage, citric acid [C6] loses a carbon as
CO2 to form α-ketoglutarate [C5] while generating one NADH molecule

✳ In the five-carbon stage, α-ketoglutarate [C5] loses a carbon as CO2 to form succinyl
coenzyme A (succinyl-CoA) [C4] while generating one NADH molecule

✳ Through a series of reactions in the four-carbon stage, succinyl-CoA [C4] is converted


to OAA [C4] while generating one NADH molecule, one FADH2 molecule, and one GTP
molecule (equivalent to ATP; produced via substrate-level phosphorylation)
Summary of TCA

Summary of TCA cycle per molecule of pyruvate


✳ 4 NADH molecules
✳ 1 FADH2 molecule
✳ 1 ATP (as GTP) molecule
✳ 3 CO2 molecules
ELECTRON TRANSPORT CHAIN
Electron Transport Chain
◆ In eukaryotes, the electron transport chain is located in the
mitochondrial inner membrane
✳ Comprised of a series of electron carriers arranged into four
complexes connected by coenzyme Q and cytochrome c
✳ Electrons flow from electron carrier to another
✳ Final electron acceptor is to O2

◆ Energy that is released during these transfers is captured as ATP


via the formation of proton and electrical gradients

◆ The ATP is made via the process of oxidative phosphorylation -


ATP formation driven by electron transfer between carriers
✳ NADH will drive the synthesis of 3 ATPs
✳ FADH2 will drive the synthesis of 2 ATPs
ETC in Prokaryotes
◆ The same principles as in eukaryotes apply to
the electron transport system in prokaryotes,
but the structural details differ as do
physiological responses
✳ Vary in cytochrome composition
✳ Electrons can enter at several different points
in the chain
✳ Chains may be branched
✳ Different branches may operate under
different environmental conditions
OXIDATIVE PHOSPHORYLATION
Oxidative Phosphorylation
◆ Several theories have been postulated for this mechanism of ATP
synthesis
◆ Most widely accepted theory is the chemiosmotic theory
✳ During electron transport in eukaryotes, protons move outward
from the mitochondrial matrix and electrons are transported
inward
✳ Results in the formation of a gradient of protons and a membrane
potential due to the unequal distribution of charges
✳ This unequal distribution of charges, known as the proton motive
force, drives the formation of ATP when protons return to the
mitochondrial matrix
✳ ATP synthesis occurs through the action of ATP synthase (F1
particle) attached to the inner mitochondrial membrane
 In prokaryotes, a similar process takes place with the F1 particle
located in the plasma membrane
ATP CONSERVATION
Yield of ATP
◆ Eukaryotes theoretically generate 36-38 total
ATP molecules per glucose molecule
catabolised
◆ Prokaryotes may produce less ATP (about 30
molecules per glucose molecule catabolised)
due to less efficient electron transport chain,
but still produces more energy in this manner
than in anaerobic respiration
ANAEROBIC RESPIRATION
Anaerobic Respiration
◆ Anaerobic respiration is an energy-yielding
process that involves the use exogenous
electrons acceptors other than O2
◆ Major electron acceptors in anaerobic
respiration include nitrate, sulfate, and CO2,
though various metals and organic compounds
also function as final electron acceptors.
Anaerobic respiration is less efficient than aerobic
respiration in the synthesis of ATP
FERMENTATION
Fermentations
◆ In the absence of aerobic and anaerobic respiration, NADH is not
oxidized by the electron transport chain due to the absence of an
external electron acceptor
◆ NADH must still be oxidized back to NAD+ or glycolysis will stop
◆ Many microbes use pyruvate or its derivatives as the electron
acceptor
◆ Various types of fermentations exist, but all have two common
themes:
✳ NADH is oxidized to NAD+
✳ Electron acceptor is often pyruvate or a derivative thereof that,
when partially , can lead to ATP production
◆ Some chemoorganoheterotrophs will only produce energy via
fermentation even though exogenous electron acceptors are
available
◆ Some obligate anaerobes use CO2 (or carbonate)
or sulfate as final electron
acceptors
✳ Methanogens are able to reduce CO2 to
methane (CH3)
✳ Sulfur bacteria reduce SO4
- to sulfide (S2 or hydrogen sulfide (H2S)
◆ In summary, anaerobic respiration is less
efficient than aerobic respiration in the synthesis
of ATP

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