CHAPTER 16

Respiration

INTRODUCTION

energy to drive live activit:. whether or animals require ALLSolar living energy organisms trapped by greenplants plants during is the ultimatetivitiesso source for the purpose. 1o Keep the life proce. energy available to living organisms cess ofin motion, the energy is derived by the oxidation of various photosyntheticnthetic productprode by the process called respiration. Respiration is a vital process which occurs in all living cells of the plant h the most actively respiring regions are growing regions like tioral and vegetative buds, germinating seedlings, stem and root apices. Roughly, respiration may ha called a process which includes the intake of oxygen and chemically bringsabo the oxidation and decomposition of organic compounds with the release ofenerey During normal conditions the process involves the liberation of carbon dioxide absorption of oxygen and conversion of potential energy into kinetic overall energy, The respiration process may be represented as CH1206+ 60, 6C0, + 6H,0 +686 kcal However, oxygen gas does not react with are directly . Water required which are added to molecules intermediate products to hydrogen atoms in the glucose degradation and intermediate products are case, the overall transferred to oxygen. In summary of the process would be that

CH,0,+6H,0 - > +60 6C0, + 12H,0 +Energy The utilization of water molecules at heading various are mechanism of respiration. steps explained under the Respiration is a () complex process which includes- absorption of oxygen, (i) conversion of carbohydrate (complex) to carbon substances), i.e. oxidation of dioxide and water (in) release of food, (simpie energy-a of and rest part which is may be lost utilised in various es in the form of vital proces v) formation of intermediate heat, )liberation of products playing (vi) loss carbon dioxide and different roles in olism, in weight of water, and metaD plants as a result of oxidation. Respiration 319

Decniration is, therefore, a reverse process of photosynthesis. The Table 16.1 mmarises the points of differences between the two processes. Table 16.1. Differences Between Respiration and Photosynthesis.

Respiration Photosynthesis 1. Oxygen is liberated in the process. is absorbed in the process. . Oxygen as a result 2. Carbon dioxide is absorbed and is fixed Carbon dioxide is evolved 2. inside to form carbon contaíning of oxidation of carbon containing compounds. compounds.

occurs and 3. Process occurs only in presence of light. 3. The process day night. for the 4. Light is essential for the process. 4. Light is not essential process. is 5. During the process, radiant energy (light 5. During the process, potential energy converted into kinetic energy. energy) is converted into potential energy materials used are and water. 6. Raw materials used are glucose and 6. Raw CO, oxygen. 7. The presence of chlorophyll is not 7. Presence of chlorophyll is necessary for necessary. photosynthesis. 8. Energy is released during the process 8. Energy is stored during the process hence hence it is an exothermic procesSs. it is an endothermic process. 9. Due to respiration the plant suffers with 9. By the process, the weight is gained. the loss of weight. 10. It is a catabolic process and includes 10. It is an anabolic process and includes the destruction of stored food. the manufacture of food. 11. Theprocess includes dehydrolysis and 11. It includes the processes like . and carboxylation. 12. During the breakdown of glucose 12. During the synthesis of one glucose molecule, 38 ATP molecules are formed. molecule, 18ATP moleculesare utilised. Significance of Respiration The respiration is an important process because () It releases energy which is consumed in various metabolic processes essential for plant life and activates division. (in It brings about the formation of other necessary compounds participating as important cell constituents. (iit) It converts insoluble food into soluble form. (iv) It liberates carbon dioxide and plays a part actively in maintaining the balance of carbon cycle in nature. stored () It converts energy (potential energy) into usable form (kinetic energy). TYPES OF RESPIRATION

has been divided into two of oxygen, respiration On the basis of the availability and the categories. in the presence of oxygen respiration. It takes place (A) Aerobic oxidised into carbon dioxide gets completely stored food (respiratory substrate) and water, i.e., + 686 kcal CH,O, +60, 6C0, + 6H,0

occurrence and is found in all plants. is of common This type of respiration in the absence of oxygen or when Anaerobic respiration. It takes place (B) cent. The stored food is incompletely concentration is less than one per oxygen are dioxide and water certain other compounds oxidised and instead of carbon common micro- of rare occurrence but among formed. This type of respiration is can be by organisms like yeasts and represented kcal CH0 2C,HOH +2C0, +56 324 Plant Physiology

Table 16.2. Difference Between Aerobic and Anaerobic Respiratio.lon. Aerobic respiration Anaeroblc respiration 1. It is common to all plants. . It is of rare occurrence. 2. It goes on throughout the lifc. 2. It occurs for a porary phase 3. Energy is liberated in larger quantity. In 3. Energy is liberated in lesser quantitv total, 38 ATP molecules/glucose molecule 2 ATP molecules are formed.quantity. Onl are formed. 4. The process is not toxic to plants. 4. It is toxic to plants. S. Oxygen is utilized during the process.5. It occurs in absence of oxygen 6. Thecarbohydrates are oxidised completely6. The carbohydrates are oxidised and are broken down into CO, and water. incompletely and ethyl alcohol and cark dioxide are formed. carbon 7. The end-products are carbondioxide and 7. The end-products are ethyl alcoholnol and an water. carbon dioxide. 8. The process takes place partly (gtyeolsis) |8. The process occurs only in the in the cytosol and partly (Krebs cycle)| cytosol. inside the mitochondria.

MECHANISM OF RESPIRATION

Respiratory substrate. The substrates which are broken down in respiration for release of energy may be carbohydrates, fats or proteins. Proteins are used up as respiratory substrate only when carbohydrates and fats are not available Blackman proposed that respiration in which carbohydrates are used as respiratory substrate are calledfloatingrespiration andif proteins are used, protoplasmicrespiration Fats are used as respiratory substrates after their hydrolysis to fatty acids and glycerol by lipase and their subsequent conversion to hexose sugars: Proteins serve as substrates after their breakdown to amino acids by proteolytic . As regards carbohydrates, not only simple hexose sugars like glucose and fructose but complex disaccharides particularly sucrose and polysaccharides such as starch, inulin and hemicelluloses are also used as respiratory substrates. During respiration, the complex substrates are broken down into simpler ones and finally CO, is liberated and water is formed. During oxidation of respiratony substrate, certain amount of energy is released. Part of this energy is trapped n the form of energy rich compounds such as ATP while the remaining part is lost in the form of heat. The energy trapped in ATP molecules can be used in variou ways (both for physical and chemical requirements). Here, the typical examgt or respiratory substrate, i.e. carbohydrate has been discussed. Oxidation ot and proteins are given separately. All complex carbohydrates are firstly converted into hexose (glucose or fruc oxidation o before actually entering into the respiratory process (Fig. 16.1). The oxidae glucose to CO, and water consisis of two distinguishable phases: (i) , and (i) Krebs cycle. In glycolysis glucose is converted into . Steps of glycouynirato are common to all kinds of respiration, hence may be called commo Respiration 325 em. The fate of metabolism pyruvic acid, however, etabfof oxygen. In;In of depends on the presence or the absence oxygen. presence oxygen, the final nand water (Krebs cycle) while degradation products are diox in carbon in absence of oxygen rbonjoxioxidedioxidei fermentation and lactic acid ethyl alcohol ana in lactic acid formation are formed. Starch UDPG Surcose+ UDPG Amylase

Mannose Glucose Fructose Starch Galactose

ATP +H,PO + +ATP ATP ATP Phospho- Hexokinase Hexokinasse Hexokinase J rylase Glucose Galactose ADP 1- 1-phosphate ADP ADP NPhosphaglucomutase Isomerase Mannose Glucose Fructose Glucose-6-phosphate 6-phosphate 6-phosphate 6-phosphate

GLYCOLYSIS before Fig. 16.1. Schematic conversion of complex carbohydrates entering into Glycolysis. acid formation processes fermentation, anaerobic respiration and lactic Glycolysis, of mitochondria while Krebs cycle occurs in the matrix occur freely in the cytoplasm in prokaryotic cells. Enzymes the eukaryotic cells and on the surface of mesosomes These in of called cytosol. found in the soluble portion cytoplasm, of glycolysis are and Such enzymes time and are required again again. remain active throughout the life are called constitutive enzymes.

GLYCOLYSIS

Common respiratory pathway, MeyerhofParanaspathway, (EMPpathway = Embden acid is termed Cytoplasmic respiration). from glucose to pyruvic degradation is also known The course of stepwise glycolytic pathway of its tracers, the a5 After the name gycolysis. (EMP Paranas pathway pathway). as Embden Meyerhof follows: represented as can be broadly Glycolysis 2C H,0,+ 4H

(Pyruvate) (Hexose) broken 6-carbon compound is which is a molecule glucose throughthrough states that a of a 3-carbon compound nus acid which is pyruvic occurs in the following two molecules of reactions. It downinto into integrated a larg stepwise closely thenumber of phosphorylated with Three important phases. molecule is the glucose this two glycolysis, structure. For phase first phase of into its thetne groups he introduction of two phosphate moleeOIeCules of ATP are needed. 326 Plant Physiolog

The second phase involves the breaking up of 6-carbo compound Fructo 1,6-diphosphate into two molecules of 3-carbon compounds,compounds, i,e 3-Phosphoglyceraldehyde and Dihydroxyacetone phosphate. These two 3-carh.e. compounds are interconvertible. The third phase involves degradation of 3-PGAld into pyruvic acid with h production of four molecules of ATP. As in the phosphorylation of glucose durin the first phase where two molecules of ATP have already been used up, thereuring up, there s reactions. of ATP during glycolytic a net gain of only two molecules detailed as follows. The various steps of glycolysis are hexokinase and with the th thethe helpheln First all in presence of of (1) of molecule is phosphorvlato one ATP molecule, the sixth carbon position ofgluc0se ted Glucose-6-phosphate. ATP is, however, converted and glucose is converted into erted as. follows. into ADP. The reaction may be represented Hexokinase Glucose Glucose-6-phosphate Mg ATP ADP

(2) Next reaction involves the isomerisation and conversion of Glucose-6-phosphate into Fructose-6-phosphate. The conversion is catalysed by the enzyme phosphoglucoisomerase Phosphoglucoisomerase Glucose-6-phosphate F 2Fructose-6-phosphate 3) Now the first carbon of Fructose-6-phosphate also gets phosphorylated with the help of another ATP molecule in the presence of the enzyme phospho- fructokinase and is converted into fructose-1,6-diphosphate. Magnesium are needed for enzymatic activity of . Phosphofructokinase Fructose-6-phosphate- >Fructose-1,6-diphosphate MgMg* ATP ADP 4) This two-fold 1, phosphorylation of hexose 6-diphosphate breaks into permits its break-up. Fru of two molecules of the enzyme 3-carbon in aldolase. The two 3-carbon compounds p3. and compoundsompounds formedformeu ar Phosphoglyceraldehyde poundsns are and an Dihydroxyacetone phosphate. Thesese two of interconvertible equilibrium is two rsion maintained between them. The versio enzyme3-Phosphoglyceraldehyde phosphotrioseisomerase. and interco bythethe Dihydroxyacetone phosphate isis catalysedcatay CH,O) CH,O(P)

CHO C O C 0 Aldolase

HOCH CH,OH HCOH Dihydroxyacetone

HCOH CH,O(P) phosphate 3-Phospho HCOH glyceraldehyde

CH,O(P) Fructose-1, 6-diphosphate CHO CH,OCP)

HCOH C O Phosphotrioseisomerase CH,O(P) CH,OH Phosphoglyceraldehyde Dihdroxyacetone phosphate Here (P) refers to phosphate, i.e., PO,H2 and the attachments next is the oxidation 3-phosphoglyceraldehyde (5) The step of 3-Diphosphoglyceric to the molecule forming 1, of inorganic phosphate H,PO, release of two molecule is oxidised with the acid. The 3-phosphoglyceraldehyde in the of the reaction are coupled and two protons (H*). The two steps electrons of3-Phosphoglyceraldehyde) the supplied by one step (oxidation sense that energy phosphate of organic linkage between inorganic is utilised by the other step (formation 1, 3- in the C, position to produce and oxidised 3-Phosphoglyceraldehyde the serve to most of energy The two in fact trap Diphosphooglyceric acid). steps heat. This would be dissipated as oxidation which otherwise simply liberated in two of the above ATP in the next step. The steps is however recovered as and energy phosphoglyceraldehyde dehydrogenase the enzyme reaction are catalysed by in released are however used up reducing the two electrons along with protons (H*) NAD to NADH + H C- O(P) CHO Phospoglyceraldehyde dehydrogenase HCOH + NADH + H* HCOH+ Pi + NADD CH,O(P) CH,O(P) 1,3-Diphosphoglyceric acid. 3-Phosphoglyceraldehyde phosphate) (Pi represents inorganic 3phosphoglyceraldehyde to 1,3-diphospho0 With the conversion of the (6) it more Dihydroxyacetone is affected and to maintain of acid a shift in the balance Syceric 3-Phosphoglyceraldehyde. PHOsphate is converted into formation of 1,3- phosphate) in the of Pi (inorganic of incorporation released in the oxidation he with the energy phosphoglyceric acid along this attaches because in the next step phosphate 3-PhTnOsphoglyceraldehyde is important 328 Plant Physiology

itself with ADP to produce ATP where 1, 3-Diphosphoglyceric acid is convertea into 3-Phosphoglyceric acid in presence of the enzymephosphogyceriu acid kinase. This transferTed from another kind of reaction in which a phosphate group is already phosphorylated compound to ADP to form ATP is called transphosphorylationsphorylation (or substrate phosphorylation). COO(P) COOH 10spoglyceric acid kinase HCOH + ATP HCOH +ADP CH,OP) CH,O(P) 1,3-Diphosphoglyceric acid 3-Phosphoglyceric acid

3-Phosphoglyceric acid (7) In presence of the enzyme phosphoglyceromutase, IS transformed to 2-phosphoglyceric acid.

COOH COOH Phospoglyceromutase HCOH HCO(P) CH,O(P) CH,OH 3-Phosphoglyceric acid 2-Phosphoglyceric acid

(8) In the next step catalysed by the enzyme one molecule of water is eliminated from 2-Phosphoglyceric acid and it is converted into 2-.

COOH COOH Enolase HCO(P) 2 Mg C-O(P) + H,O CH,OH 2-Phosphoglyceric acid CH, 2-Phosphoenolpyruvic acid 9) The removal of water from acid structure and changes its internal 2-Phosphoglyceric alters its molecular part of the distribution in such a molecule energy is concentrated way that a much greater When this in the region of the phosphate group is taken over phosphate group: acid by ADP in the acidphosphoenolpyruvic to pyruvic acid in conversion o kinase, a considerable the presence of the Fig. part of energy is enzyme pyriv 16.3). This kind of conserved as ATP reaction is also called (For details s transphosphorylation regctio Resplration 329 GLYCOLYSIS Glucose Heokinase Mg** ATP ADP Glucose-6-phosphate Phosphoglucoisomerase Fructose-6-phosphate Phosphofructokinase - ATP Mgt

AdolaseADP-Fructose-1,6-diphosphate 2Pi Diphydroxy acetone phosphate Phosphotriose 3-Phosphoglyceraldehyde isomerase 2PI Phosphoglyceraldehyde +2NAD dehydrogenase 2NADH+H*-1,3-Diphosphoglyceric acid (2 molecules) Phosphoglycerokinase Each molecule -2ADP 2ATP-3-Phosphoglyceric acid Phosphoglyceromutase

2-Phosphoglyceric acid

Enolase

2-Phosphoenolpyruvic acid

Pyruvic acid kinase 2ADP

2ATP Pyruvic acid

CO,+Ethyl alcohol Entrance into Fermentation Krebs cycle of glycolysis. Fig. 16.2. Schematic representation