CHAPTER 12: RESPIRATION IN PLANTS

 The breaking of the C – C single bond of complex compounds through oxidation

within the cells, leading to the release of a considerable amount of energy and the
trapping of this energy for the synthesis of ATP, is called cellular respiration.
 During cellular respiration or oxidation within a cell, all the energy contained in
compounds is not released in a single step because in single if the complete energy is
released it may be converted into heat. So, the energy during cellular respiration is
released in a series of slow stepwise reactions controlled by enzymes, and it is trapped
as chemical energy in the form of ATP.
 ATP, thus synthesised, is broken down whenever and wherever energy needs to be
utilised; hence, ATP acts as the energy currency of the cell.
 The compounds that are oxidised during the process of cellular respiration are called
respiratory substrates; these are usually carbohydrates, but fats, proteins & organic
acids can also be used as respiratory substrates.
 Primarily, carbohydrates (mainly glucose) are used as respiratory substrates. In the
absence or less availability of carbohydrates, the respiratory substrates can be fats &
proteins.

 The sequence of use of respiratory substrates – (i) Carbohydrate, (ii) Fat, (iii) Protein.
 Organic acids such as malic acid, etc., can be used as respiratory substrates in some
plants under certain conditions.

FEATURES OF CELLULAR RESPIRATION

i. Cellular respiration is an amphibolic process. The carbon skeleton (Intermediates of

respiration) produced during respiration is used as a precursor for the biosynthesis of
other molecules in the cell.
ii. Cellular respiration is an exergonic process. The breaking of C–C bonds of complex
compounds through oxidation within the cells leads to the release of a considerable
amount of energy.

12.1 DO PLANTS BREATHE?

 Plants, unlike animals, have no specialised organs for gaseous exchange but they have
stomata and lenticels for this purpose.
 There are several reasons why plants can get along without respiratory organs:
(a) First, each plant part takes care of its own gas exchange needs. There is very little
transport of gases from one plant part to another.
(b) Second, plants do not present great demands for gas exchange. Roots, stems and
leaves respire at rates far lower than animals do. Only during photosynthesis are large
volumes of gases exchanged, and each leaf is well adapted to take care of its own
needs during these periods. When cells photosynthesise, the availability of O 2 is not a
problem in these cells since O2 is released within the cell.
(c) Third, the distance that gases must diffuse even in large, bulky plants is not great. In
stems, the ‘living’ cells are organised in thin layers inside and beneath the bark. They
also have openings called lenticels. The cells in the interior are dead and provide only
mechanical support. Thus, most cells of a plant have at least a part of their surface in
contact with air. This is also facilitated by the loose packing of parenchyma cells in
leaves, stems and roots, which provide an interconnected network of air spaces.

TYPES OF RESPIRATION
(A) On the basis of type of respiratory substrates:
(1) Floating respiration: - When carbohydrates or fats are oxidised inside the cell.
Carbohydrates and fats are floating inclusions of cells. Thus, this is called floating
respirations.
(2) Protoplasmic respiration: - When protein is oxidised inside the cell. This occurs in
starved cells. Protein is a constituent of protoplasm; thus, this is called protoplasmic
respiration.
(B) On the basis of the presence or absence of O2:
(1) Aerobic respiration
(2) Anaerobic respiration/Fermentation
In eukaryotic cells, aerobic respiration is completed in four stages:
Stage 1: Glycolysis
Stage 2: Oxidative phosphorylation
Stage 3: Kreb’s cycle
Stage 4: Oxidative phosphorylation
12.2 GLYCOLYSIS

 The term glycolysis has originated from the Greek words glycos for sugar, lysis means
splitting.
 The scheme of glycolysis was given by Gustav Embden, Otto Meyerhof & J. Parnas
so glycolysis is often called EMP pathway.
 In anaerobic organisms, it is the only process in respiration.
 Glycolysis is the common phase between aerobic and anaerobic respiration.
 Glycolysis occurs in the cytoplasm of the cell.
 Glycolysis is defined as the process of partial oxidation of glucose to form two
molecules of pyruvic acid.
 In plants, glucose is derived from sucrose (the end product of photosynthesis) or from
storage carbohydrates.
 Sucrose is converted into glucose and fructose by the enzyme invertase. These 2
monosaccharides enter the glycolytic pathway.

MECHANISM OF GLYCOLYSIS

Step 1: Glucose is phosphorylated to glucose-6-

phosphate by ATP in the presence of enzyme
hexokinase.

Step 2: Glucose-6-phosphate is isomerised into

fructose-6-phosphate.

Step 3: Fructose-6-phosphate is further

phosphorylated by means of ATP to fructose-
1,6-bisphosphate. It is an irreversible reaction
and a rate-limiting step in glycolysis.

Step 4: Fructose-1,6-bisphosphate is split into

one molecule of 3C compounds
glyceraldehyde-3-phosphate/3-
phosphoglyceraldehyde (3-PGA) and
dihydroxyacetone phosphate (DHAP).

Step 5: DHAP is then isomerised to 3-PGA.

Thus, 2 molecules of 3-PGA are formed from
a single fructose-1,6-bisphosphate.

Step 6: Glyceraldehyde-3-phosphate loses 2 H-atoms to NAD to form NADH2 and

accepts inorganic phosphate to form 1,3-bisphosphoglyceric acid (BPGA).

Step 7: BPGA loses a phosphate in 1 st carbon and is converted to 3-phosphoglyceric

acid (PGA). The lost inorganic phosphate is accepted by an ADP to form an ATP.

Step 8: 3-phosphoglycerate (3-PGA) is changed into its isomer 2-phosphoglycerate (2-

PGA) by transfer of phosphate from 3rd to 2nd carbon of 3-PGA.
Step 9: 2-PGA (2-phosphoglyceric acid) loses an H 2O molecule to form phosphoenol
pyruvate (PEP).

Step 10: PEP loses a phosphate and is converted into pyruvic acid or pyruvate. The
loosed inorganic phosphate is accepted by an ADP to form an ATP.

Product of glycolysis:
2 molecules of pyruvic acid, net gain of 2ATP molecules, 2 molecules of NADH 2

Fate of Pyruvic acid:

12.3 ANAEROBIC RESPIRATION OR FERMENTATION

Fermentation is a kind of anaerobic respiration carried out primarily by fungi and

bacteria.

Types of fermentation:

1. Alcoholic fermentation
 Pyruvic acid formed at the end of glycolysis is converted to alcohol by using two
enzymes, pyruvic acid decarboxylase and alcohol dehydrogenase.
 In the first step, pyruvic acid is decarboxylated, resulting in the formation of
acetaldehyde and CO2.

 In the second step, acetaldehyde is reduced to alcohol by 2NADH + H+

Example: This is commonly seen in yeast and certain bacteria.

Ethyl alcohol does not stay inside micro-organisms but is excreted. Accumulation of
alcohol beyond a certain limit can, however, kill the microorganisms. Yeasts poison
themselves to death when the concentration of alcohol reaches about 13%. Therefore,
the maximum concentration of alcohol in beverages that are naturally fermented is 13 %.
A higher concentration of alcohol in a beverage is achieved through distillation.

2. Lactic acid fermentation

Pyruvic acid formed at the end of glycolysis is reduced to lactic acid by

homofermentative lactic acid bacteria (Lactobacillus lacti).

During vigorous exercise, when oxygen is inadequate for cellular respiration, pyruvic
acid is reduced to lactic acid by using the enzyme lactate dehydrogenase.
 Fig. Major pathways of anaerobic respiration

Special features of alcohol and lactic acid fermentation:

 In both lactic acid and alcohol fermentation, not much energy is released; less than
seven per cent of the energy in glucose is released, and not all of it is trapped as high
energy bonds of ATP.
 Both processes are hazardous - either acid or alcohol is produced.
 The net gain in both types of fermentation is 2ATP.
 Alcohol fermentation results in the release of CO 2 along with ethanol while lactic acid
fermentation releases lactic acid only.
+¿¿ + ¿¿
 The reducing agent is NADH + H which is re-oxidised to NAD in both processes.

12.4 AEROBIC RESPIRATION

 It is the process by which organisms can carry out complete oxidation of glucose and
extract the energy stored to synthesise a larger number of ATP molecules needed for
cellular metabolism. In eukaryotes, these steps take place within the mitochondria,
and this requires O2.
 The aerobic respiration process leads to the complete oxidation of organic substances
in the presence of oxygen and releases CO 2, water and a large amount of energy
present in the substrate. This type of respiration is most common in higher organisms.
  For aerobic respiration to take place within the mitochondria, the pyruvic acid formed
during glycolysis is transported from the cytoplasm into the mitochondria.
 The crucial events in aerobic respiration are:
i. The complete oxidation of pyruvate by the stepwise removal of all the hydrogen
atoms, leaving three molecules of CO 2. It occurs in two steps:
(a) Formation of Acetyl coenzyme A.
(b) TCA cycle (Tricarboxylic acid cycle) or Krebs cycle or Citric acid cycle.

This process takes place in the matrix of mitochondria.

ii. The passing on of the electrons removed as part of the hydrogen atoms to molecular
O 2 with simultaneous synthesis of ATP through ETS.

This process takes place on the inner membrane of the mitochondria.

(a) Formation of Acetyl CoA or Oxidative phosphorylation:

 For this, the pyruvate (final product of glycolysis) is transported from the cytoplasm
into the mitochondrial matrix.
 Pyruvate undergoes oxidative decarboxylation to form acetyl CoA in the presence
of pyruvate dehydrogenase.
 It requires coenzymes NAD+ and coenzyme A.
 During this process, 2 molecules of NADH+H+ are produced from 2 molecules of
pyruvic acid.
 The acetyl CoA then enters the tricarboxylic acid cycle (TCA cycle).
 This step is called the gateway step or link reaction because acetyl CoA acts as a
connecting link between glycolysis and Kreb’s cycle.
(b) TCA cycle (Tricarboxylic acid cycle) or Kreb’s cycle or Citric acid cycle:
 This cycle was discovered by Hans
Krebs, which won him a Nobel
Prize.
 It occurs in the mitochondrial
matrix.
 This cycle is called the citric acid cycle because of the formation of citric acid in the
first step of this cycle.
 It is also known as the TCA cycle because many intermediate compounds formed in
the cycle have three carboxyl groups.
 The reactions of the cycle require the presence of oxygen and are confined to the
mitochondrial matrix.
 This cycle serves as a common oxidative pathway for carbohydrates, fats and proteins.
 All enzymes are soluble in the mitochondrial matrix, but succinate dehydrogenase is
found attached to the inner mitochondrial membrane.
 During Kreb’s cycle, acetyl CoA is completely oxidised into CO2.

MECHANISM OF TCA CYCLE:

1) Acetyl CoA (2C) combines with oxaloacetic acid (OAA – 4C) and water to form
citric acid (6C) in the presence of the enzyme citrate synthase. It is the first product
of Kreb’s cycle. During this, a molecule of CoA is released.
2) Citrate is isomerised into isocitrate.
3) Isocitrate is decarboxylated to form α – ketoglutaric acid (5C). This step reduces
NAD+ to NADH+H+.
4) α – ketoglutaric acid is decarboxylated to form succinyl-CoA (4C). This step reduces
NAD+ to NADH+H+.
5) Succinyl-CoA is converted to succinic acid. This step leads to a substrate-level
phosphorylation of GDP to form GTP. This reaction is simultaneously coupled with
the synthesis of ATP from ADP.
6) Fumaric acid is formed in the next step from succinic acid, which is catalysed by
succinate dehydrogenase. During this reaction, the FAD molecule is reduced to
FADH2.
7) Malic acid is formed by the action of fumarase on fumaric acid. This step involves the
addition of water molecules.
8) Regeneration of oxaloacetic acid is the last step catalysed by malic dehydrogenase,
which reduces NAD+ to NADH+H+ by removing hydrogen from malic acid.
 One molecule of ATP (via direct GTP), three NADH 2, one FADH 2 and two
molecules of CO 2 are released per molecule of acetyl CoA oxidised.
 However, as two molecules of pyruvic acid are formed from one glucose
molecule, TCA cycle must occur twice for each molecule of glucose respired.
 Therefore, 2ATP, 6 NADH 2 and 2 FADH 2 are formed from 2 molecules of acetyl
CoA (coming from one molecule of glucose).

The summary equation for this phase of respiration (Oxidative decarboxylation + Kreb’s
cycle) may be written as follows:

Difference between Glycolysis and Krebs cycle

(c) Electron Transport System (ETS) and Oxidative Phosphorylation

 The metabolic pathway through which the
electron passes from one carrier to another
is called the electron transport system
(ETS).
 This process occurs in the inner
mitochondrial membrane.
 During this process, hydrogen acceptors
get re-oxidised, and ATP is produced.
Components of ETS and ATP synthase

Name of complexes Components

of ETS

Complex I (NADH FMN, Fe-S

dehydrogenase complex)

Complex II (Succinic FAS, Fe-S

dehydrogenase complex)

Complex III (Cytochrome Cyt b, Cyt c1,

C reductase complex) Fe -S

Complex IV (Cytochrome Cyt a, Cyt a3,

C oxidase complex) 2Cu centres

Complex V (ATP F0 (integral)

synthase/ATPase/Oxysome) – F1
(peripheral)

MECHANISM OF ETS:

1. Electrons from NADH produced in the mitochondrial matrix during citric acid cycle
are oxidised by an NADH dehydrogenase (complex I), and electrons are then
transferred to ubiquinone located within the inner membrane.
2. Ubiquinone (UQ) receives reducing equivalents through FADH 2 (complex II) that is
generated during the oxidation of succinate in the citric acid cycle.
3. The reduced ubiquinone (ubiquinol) is then oxidised with the transfer of electrons to
cytochrome c via cytochrome bc1 complex (complex III).
Cytochrome c is a small protein attached to the outer surface of the inner membrane
and acts as a mobile electron carrier for the transfer of electrons between complex III
and IV.
4. Cytochrome c oxidase (Complex IV – complex containing cytochrome a and a 3 and
2Cu centres) pass electrons onto O 2 which is reduced to H2O by the addition of H+.
This is called terminal oxidation.

Oxidative phosphorylation
 i. When the electrons pass from one carrier to another via complex I to IV in the
electron transport chain, they are coupled to ATP synthase (complex V ) for the
production of ATP from ADP and inorganic phosphate.
ii. The proton gradient or proton motive force (PMF) required for phosphorylation is
obtained with the use of the energy of oxidation-reduction. Therefore, it is called
oxidative phosphorylation. The mechanism of membrane-linked ATP synthesis is
explained by the chemiosmotic hypothesis given by Peter Mitchell.
iii. The inner mitochondrial membrane is permeable to protons only in the region of
F 0−F 1 or elementary particles or ATP synthase.
iv. The energy released during the electron transport system is utilised in synthesising
ATP with the help of ATP synthase (complex V ). This complex consists of two major
components, F 1 and F 0.
v. The F 1 headpiece is a peripheral membrane protein complex and contains the site for
synthesis of ATP from ADP and inorganic phosphate. F 0 is an integral membrane
protein complex that forms the channel through which protons cross the inner
membrane.
vi. The passage of protons through the channel is coupled to the catalytic site of the F 1
component for the production of ATP. For each ATP produced, 2 H +¿¿pass through F 0
from the intermembrane space to the matrix down the electrochemical proton
gradient.

Three pairs of protons are pushed during oxidation of each NADH + H +¿¿ and two pairs of
protons are pushed during oxidation of each FADH 2.
Complete oxidation of NADH forms 3ATP molecules, while one FADH 2 forms 2ATP
molecules.
12.5 RESPIRATORY BALANCE SHEET
Theoretical energy calculation for complete oxidation of one glucose molecule

Stage of Number ATP synthesis ATP gain ATP Net

respiration of turns through through consumed gain
substrate level oxidative
phosphorylation phosphorylation
Glycolysis 1 4 6 or 4 2 8 or 6
Link 2 0 6 0 6
reaction
Krebs cycle 2 2 22 0 24
Total - 6 34 or 32 2 38 or
36

It is possible to make calculations of the net gain of ATP for every glucose molecule
oxidised, but in reality, this can remain only a theoretical exercise. These calculations can
be made only on certain assumptions that:

(a) There is a sequential, orderly pathway functioning, with one substrate forming the
next and with glycolysis, TCA cycle and ETS pathway following one after another.
(b) The NADH, synthesised in glycolysis is transferred into the mitochondria and
undergoes oxidative phosphorylation.
(c) None of the intermediates in the pathway are utilised to synthesise any other
compound.
(d) Only glucose is being respired - no other alternative substrates are entering the
pathway at any of the intermediary stages.

Total net gain of ATP=36 or 38 depending upon the type of shuttle system used in aerobic
respiration. In most eukaryotic cells the net gain of ATP is 36 molecules.

But, these kinds of assumption are not really valid in a living system; all pathways work
simultaneously and do not take place one after another; substrates enter the pathways and
are withdrawn from it as and when necessary; ATP is utilized as and when needed;
enzymatic rates are controlled by multiple means.

Yet, it is useful to do this exercise to appreciate the beauty and efficiency of the living
system in extraction and storing energy. Hence, there can be a net gain of 36 ATP
molecules during aerobic respiration of one molecule of glucose.

12.6 AMPHIBOLIC PATHWAY

 Glucose is the favoured substrate for respiration.

 All carbohydrates are usually first converted into glucose before they are used for
respiration.
 Other substrates can also be respired, but then they do not enter the respiratory
pathway at the first step.
 Fats are first broken down into glycerol and fatty acids. Fatty acids are degraded into
acetyl CoA and enter the pathway.
 Glycerol would enter the pathway after being converted to PGAL
(phosphoglyceraldehyde).
 Proteins are degraded by proteases into amino acids and, depending on their structure,
enter the pathway at some stage within the Krebs cycle or even as pyruvate or acetyl
CoA.
 Breaking down processes within the living organism is catabolism and synthesis of
molecules within the living organism is anabolism.
 The respiratory pathway is mainly a catabolic process that serves to run the living
system by providing energy.
 The pathway produces a number of intermediates. These intermediates of the pathway
are precursors of various compounds e.g.,
A. Acetyl CoA= raw material for carotenoids, terpenes, gibberellins etc.
B. Succinyl CoA= raw material for chlorophyll, cytochrome.
C. Oxaloacetic acid= raw material for alkaloids, pyrimidines.
D. α-ketoglutaric acid= raw material for amino acid synthesis.
 Fig. Interrelationship among metabolic pathways showing respiration mediated
breakdown of different organic molecules to CO2 and H2O

Thus, the respiratory pathway is involved in both anabolism and catabolism, it would
hence be better to consider the respiratory pathway as an amphibolic pathway rather than
as a catabolic one.

12.7 RESPIRATORY QUOTIENT

The ratio of volume of CO2 evolved to the volume of O2 consumed in respiration is called
the respiratory quotient (RQ)

The respiratory quotient depends upon the type of respiratory substrate used during
respiration.

a) When carbohydrates are used as substrate and are completely oxidised, the RQ will be
1, because equal amounts of O2 and CO2 are consumed and evolved, respectively, as
shown in the equation below:
b) When fats are used in respiration, the RQ value is less than 1. Calculations for a fatty
acid, tripalmitin, if used as a substrate is shown:

c) When proteins are respiratory substrates, the ration would be 0.9

d) RQ value for oxalic acid is 4 and 1.33 for malic acid.