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  • 3.1.10. Biosynthesis of starch and sucrose
  • 3.2. Photosynthesis: Physiological and Ecological Considerations
  • 3.3.3.T.C.A. CYCLE/KREB’S CYCLE:
  • 3.4.3. SYNTHESIS OF MEMBRANE LIPIDS Biological membrane
  • 3.4.6. SUM UP:-
  • 3.4.7. ASSIGNMENT
  • 3.4.8. REFERENCE


Sc BOTANY Self Instructional Material

M.Sc – Previous PAPER-IV Plant physiology and metabolism UNIT-III Block II

Madhya Pradesh Bhoj (Open) University BHOPAL



Editor: Dr. (Smt.) Renu Mishra HOD, Botany & Microbiology Sri Sathya Sai College for Women, Bhopal Writer: Smt. Shikha Mandloi Asst. Prof. Microbiology Sri Sathya Sai College for Women, Bhopal



AND LIPID METABOLISM 3.0 Introduction: - PHOTOCHEMISTRY & PHOTOSYNTHESIS. 3.1.1. Objectives 3.1.2. Genral Concepts & Historical background. 3.1.3. Evolution of photosynthetic apparatus. 3.1.4. Photosynthetic pigments and light harvesting complexes. 3.1.5. photo-oxidation of water, & electron transport chain. 3.1.6. Calvin cycle for carbon assimilation. 3.1.7. Photorespiration and its significance. 3.1.8. The C4 cycle. 3.1.9. CAM Pathway. 3.1.10. Biosynthesis of Starch and Sucrose. 3.2. 3.3 Physiological and ecological consideration. RESPIRATION.

3.3.1. Overview of plant respiration. 3.3.2. Glycolysis. 3.3.3. The TCA cycle. 3.3.4. Electron transport chain and ATP synthesis. 3.3.5. Pentose phosphate pathway. 3.3.6. Glyoxylate cycle. 3.3.7. Alternative oxidase system. 3.4. LIPID METABOLISM. 3.4.1. Structure and function of lipids. 3.4.2. Fatty acid biosynthesis. 3.4.2. Synthesis of membrane Lipids. 3.4.3. Structural lipids & storage lipids. 3.4.4. Lipids catabolism. 3

Because of the production of energy rich substances in the presence of light by chlorophyll.5.7 References 3. 4 C6H12O6 + 6H2 O + 6O2 .3. 3.0. You can’t imagine life on earth without plants.The word photos means light and synthesis means putting together. this process is called photosynthesis. And thus.6.a carbohydrate compound glucose is produced and stored as starch.5.6 Assignments 3.  You will know about important. & quantasome.6.Thus. intermediate compounds and enzymes involved in the process. respiration and lipid metabolism.  How does electron transport chain works.  Why photosynthesis process is studied as dark reaction and light reaction. When we say plant synthesize.1. Let us sum 3.OBJECTIVES: After learning this unit you should be able to understand the  Mechanism of conversion of solar energy into chemical energy. As you know plants are the most significant living being on earth which produce O2 and harvest solar energy for others. in the anabolic step of metabolism. the formation of carbohydrates from CO2 and water by illuminated green cells is called as photosynthesis.cal of energy is stored in chemical form. INTRODUCTION In this unit you will learn about plant physiology related to photosynthesis. their own food it is carbohydrate. when process occurs in nature in a continues manner.1.  Basic structure and functional organization of chloroplast. In other words photosynthesis is a process in which carbon dioxide is converted into carbohydrates in the presence of water and chlorophyll by all organisms containing chlorophyll Light 6CO2 + 12HO2O Chlorophyll As you know it is during this process 686 K.

Park and beggins (1964) called photosynthetic units present in granum discs quantasome. 5.1. 3.g. 3. SIGNIFICANCE OF PHOTOSYNTHESIS TO MANKIND 1. resins. Role of water as reducing agent. Photosynthesis decreases the concentration of CO2 which is being added to the atmosphere by the process of respiration of living beings and burning of organic fuels. timber. 3.2.BGA and bacteria have photosynthetic lamellae as photosynthetic apparatus. Other developments are listed in tabular form as follows. It provides food either directly as vegetable or indirectly as meat or milk of animals which in turn are fed on plants. for the first time pointed out role of sunlight in photosynthesis. 2.1. All useful plant products are derived from the process of photosynthesis e.3. fiber. STRUCTURE 5 . oils. It maintains the equilibrium of O2 and CO2 in the atmosphere. 4. Life on earth is possible because of photosynthesis. Hales in1927. These are the ultimate sites of photosynthesis. etc. rubber. PHOTOSYNTHETIC APPARATUS: In plants chloroplast are the organelles involved with photosynthesis process. GENERAL CONCEPTS & HISTORICAL BACK GROUND The process was unknown till 17 th centaury.

Thus. proteins. known as thylakoids. Each granum is composed of about 10 to 50 disc like superimposed membranous structures. The size of grana varies from 0. These interconnecting membranes of the grana are known as the stroma lamellae or fret. chloroplast is said to be a “semiautonomous cell organelle”. 2. In a granum these thylakoids are arranged in parallels to form the stakes. Limiting membrane: chloroplast is bounded by double membraned lipoprotein a covering. About 40-60 Å thick. 6 . Each thylakoid is separated from the stroma or the matrix of the chloroplast by its unit membrane. Stroma contains several small cylindrical Structures which are called grana. The grana of the chloroplast are interconnected by tubules given out by the membranes of certain thylakoids into the matrix. Each granum contains chlorophyll inside it. Chloroplast also contains 70S ribosomes. Stroma or matrix: The stroma is the inner matrix of the chloroplast which fills the inner hollow space.7 µ. Photosynthesis: The main function of chloroplast is to synthesize organic food materials by the process of photosynthesis. Photosynthesis takes place in grana. A chloroplast may contain 40 to 60 grana in the matrix. (ii) Dark reaction: Takes place in stroma.Chloroplast Ultra structure: Electron microscopic studies reveal that chloroplast is composed of following two parts. (i) Light reaction: Takes place in grana. due to the presence of above mentioned substances along with its genetic material.3 to 1.Each chloroplast contains lipids. It contains starch granules and osmophilic droplets. DNA and RNA all the pigments and electron carriers related with photosynthesis is present in the thylakoid. Functions of chloroplast: 1. 1.

There are three types of pigments in photosynthetic cells: 1. Euglenophyceae higher algae plants. Chlorophylls.No. Purple sulphur bacteria. During photosynthesis chloroplast absorb CO2 from atmosphere and photolyse H2O to release O2. This O2 is utilized by living beings during respiration. 7. Green sulphur bacteria. During light reaction of photosynthesis. This ATP and NADPH2 are required for the reduction of CO2 during dark reaction of photosynthesis. Red algae (Rhodophyta). and Brown bacteria. Chlorophylls are found within specialized structures called chloroplast.. 1. phyll=leaf): Chlorophyll is a green pigment. S. algae PIGMENTS AND LIGHT HARVESTING (Phaeophyceae). Diatoms and pyrrophyta. Chlorophylls and carotenoids are insoluble in water. Chlorophyll (G. and Mg. 3. while phycobilins are found within phycobilisomes. 3. phosphorylation of ADP takes place.2.phycobilins.1. 2. 4. 3. chlor=green. Golden Bacteriochlorophyll C55H74O6N4Mg Bacterioviridin (Xanthophyceae). 2. It is made up of 5 types of elements C. 6. N.K.Carotenoids and 3. chloroplast controls the concentration of O2 and CO2 in the atmosphere.4PHOTOSYNTHETIC COMPLEXES Chloroplast or chromatophores contain pigments which convert light energy into chemical energy during photosynthesis. algae 7 . which results ATP generation. Found within chloroplast of all green plants and in involved in photosynthesis. O. Thus. Type of chlorophyll Chemical formula Chlorophyll-a C55H72O5N4Mg Chlorophyll-b Chlorophyll-c Chlorophyll-d Chlorophyll-e C55H70O6N4Mg C35H32O5N4Mg C54H70O6N4Mg Not fully known Distribution All green plants except photosynthetic green (chlorophyceae). H. 5.

First of all plants absorb light energy with the help of their pigment systems. *Chl-a and b differ because in Chl-b there is a–CHO group instead of a-CH3 group at third carbon atom in II pyrrole ring. Phycoerythrin. The centre of tertapyrrole is occupied by bivalent magnesium (Mg++) which is complexed with nitrogen atoms of four pyrrole rings. It absorbs light energy in the mid region of visible spectrum and transfer their absorbed energy to chlorophyll molecules. (i) (ii) (iii) Head: It is made up of pyrrole group. Stoll and Fischer (1912) described the structure of chlorophyll molecule for the first time. 8 .Porphyrin ring consists of four pyrrole rings. Joined together by methane bridge (-CH3 bridge). They pick up nascent O2 released during photo oxidation of water and change them into molecular state. (Tetrapyrrole).are red or blue coloured pigments bound in BGA. they protect the chlorophyll molecules from photo-oxidation. Photosynthesis is a multistep oxidation-reduction reaction. Pyrrole head: chlorophyll is a magnesium porphyrin compound.5MECHANISM OF PHOTOSYNTHESIS PHOTO-OXIDATION OF WATER AND ETC. viz. phycocyanin. According to modern scientists. 2. The phytol chain is responsible for lipoidal solubility of the chlorophyll. (iv) Phytol tail: It is hydrophobic in nature and made up of alcohol phytol (C20H39OH). Carotenoids: It is yellow or orange colored pigment usually found in close association with chlorophylls.1. According to him chlorophyll is made up of two parts like that of a tadpole larva. Tail: It is made up of phytol group. the following three processes will take place during photosynthesis: 1. It is hydrophilic in nature.Structure of chlorophyll: Will Statter. 3. Thus. They occur in thylakoids and act as accessory pigment of photosynthesis. Phycobilins:. 3.

These antenna chlorophyll absorb light energy and transfer them into photoreaction centre or energy trapping centre. then they were going to higher energy state (excited or 9 . In this reaction light energy is utilized and formation of ATP and reducing power (NADPH + H+) takes place. 3.2. Then absorbed light energy is converted into chemical energy. The electrons required for the conversion of NADP+ into NADPH comes from water. 2. Dark reaction or Blackmann’s reaction. In PS-I energy trapping centre is P700 whereas in PS-II it is P 680. whereas third one is very complex process which does not require light hence called as dark reaction. Activation of chlorophyll-a molecules by photons of light energy: Normally chlorophyll molecule exists in ground state (or low energy state). Finally synthesis of carbohydrates takes place. Thus photosynthesis consists of two successive series of reactions: 1. Absorption of light energy by chloroplast : During photosynthesis first of all different kinds of chlorophyll molecules of leaves absorb light of different wavelengths of visible part (between 360nm to 810nm) of the spectrum and transfer it towards reaction centre of the pigment systems. Light reaction was discovered by Robert Hill (1937) hence it is also known to be as Hill reaction. Steps of Light Reaction 1. 3. in this process water functions as electron donor. This NADPH + H+ is the reduced part of redox system NADP+/NADPH. but when these chlorophyll molecules (photoreaction centre) – P 700 or P 680 absorb a photon (quantum) of light. Of the above three processes. Thus. Light or Hill reaction 2. LIGHT REACTION Light reaction takes place in grana of chloroplast and it requires light hence it is called light reaction. Transfer of light energy from accessory pigment to chlorophyll-a: All the photosynthetic pigments other than Chl-a are called as antenna or accessory pigments. first two takes place in the presence of light hence it is called as light reaction.

According to Von Niel and Frank (1941) excited molecules of chlorophylla react with water. Due to undergoing excited state energy is also comes in the outer orbit hence these are in the excited or high energy state. are either assumed in reducing NADP+ to NADPH + H+ or cycled back.S) of photosynthesis. It is believed that photolysis of water takes place due to presence of a strong oxidant which is not yet identified.4e4OH 4 H+ + 2A + 4e- 5. At this state they release electrons (e).singlet state).ions. Light Chlorophyll – a (Ground State) Chlorophyll – a Chlorophyll – a (Excited State) (Chlorophyll – a)+ + e- 4.. Photolysis or photochemical oxidation of water and evolution of oxygen: The photolysis of water molecules takes place in pigment system II in presence of Mn ++ and CI. These excited electrons are then trapped by different electron acceptors due to which chlorophyll molecule become positively charged. Chlorophyll molecule is unstable during this state. OH. And named as ‘Z’. In this state PS-II become activated and water molecules (H2O) dissociated to form H+ and OH. The extra light energy is used in the formation 10 . This process is known photochemical breakdown or photolysis of water.T. These electrons jumps from their normal orbit to high energy orbit.ions releases electrons (e-) and finally a molecule of water is formed and O2 gas is liberated. Light 4H2O Chlorophyll 4H + + 4OH4OH 2H2O + O2 2AH2 4OH.ions. Electron transport and the production of assimilatory power (NADPH + H+ and ATP) : The electron expels from P 680 and P 700 after travelling through Electron Transport System (E.

C.(Cyt-f) and plastocyanin (PC). The electrons then travel downhill and fall back to +4eV in a dark reaction through a series of PS-I. photophosphorylation or E. When a quantum of light of wavelength above 680nm is received by a molecule of PS-I the energy is transferred to a chain of other chlorophyll molecules by induction resonance. which becomes excited and releases an electron..of ATP molecule s at different place during its transport.scheme to explain the process of photophosphorylation. (i) Non-cyclic photophosphorylation : Hill and Bendal (1960) and Robinowitch and Govindjee (1965) have proposed Z. one electron is lost it doesn’t enter into PS-II. whereas in non-cyclic photophosphorylation. FRS further reduces an iron containing protein called ferredoxin. The electron from reduced ferrredoxin then reduces NADP to NADPH with the help of H+ released from H2O. until finally it is transferred to a molecule of P 700. involves the following two processes : i. When a quantum of wavelengthof light of lower wavelength is received by PS-II its reaction canter P 680 loses electron to a substance which is probably a quinine. Cyclic Photophosphorylation. Robert Hill have been first time stated that just like that of mitochondria. In cyclic photophosphorylation the electrons lost by PS-I is cycled back to it. cytochrome-f. chloroplast also utilize cytochrome. Photophosphorylation According to Arnon and associates. ii. The energy released in the transfer of electron from PQ to Cytochrome-f is utilized to convert ADP and inorganic phosphate into ATP. The electron thus does not complete the cycle as it starts from PS-II and is drained off in the carbohydrates produced by CO2 reduction.T. The carriers are cytochrome-b (Cyt-b). The ATP synthesis resulting from this type of non-cyclic 11 . According to him during light reaction. both the photochemical processes (PS-I and PS-II) takes place in a series and the product of one reaction is used in the second reaction. plastoquinone (PQ). The electron is then transferred to ferredoxin reducing substance (FRS). thus it involves both PS-I and PS-II. These electrons are accepted by ‘X’(OX))oxidised due to which it become reduced (Xred). Non-cyclic photophosphorylation. It is called as photosynthetic phosphorylation.

electron transport chain is known as non-cyclic photophosphorylation. Water molecule is utilized as a source of electron (H2 donor) in this system at the same time water molecules become dissociated into H+ and OH- ions. 4H2O 4H+ + 4OH-


4OH + 4e-

OH- ions transfer their electrons (e-) to ‘Z’ (an unknown substance) and OH radical is formed. These electrons are then transferred to PS-II and OH radical become dissociated form H2O and O2 4OH 2H2O + O2

H+ ions originated from hydrolysis of water reduces NADP+ into NADPH +H+. This NADP + H+ functions as reducing agent. Thus, we observe that the electrons released from PS-II does not again enter to PS-II hence, it is called non-cyclic photophosphorylation. 2NADP + 4H+ + 4e2NADPH + 2H+

In this process, two molecules of ATP are formed per two molecules of NADP reduced or one more molecule of oxygen evolved or two molecules of water oxidized. 2ADP + 2Pi + 2NADP+ + 2H2O 2ATP + 2NADPH + 2H+ + O2



Scheme of electron transport chain (ii) Cyclic Photophosphorylation: the cyclic photophosphorylation take place under certain condition e.g., when the amount of available NADP is low or PS-II is absent. It involves PS-I and therefore, photolysis of water and the consequent evolution of O2 does not take place. Non-cyclic electron transfer does not take place and NADPH is not formed. The electron lost by P 700 is cycled back to it through X, FRS, FD and cytochrome-b6, cytochrome –f and plastocynanin. 2ATP molecules are synthesized from 2ADP and inorganic phosphate when electron is transferred from cytochrome–b6 to PQ and from cytochrome-b to cytochrome-f . *Thus, from the above description it is clear that photochemical reaction takes palce during light reaction results: (i) Photolysis of water and release of O2 (ii) Formation of 3 ATP (iii) Formation of 2NADPH2 ATP and NADPH are used in the reduction of CO 2 during dark reaction. Similarly ATP and NADPH2 function as carrier of energy of sunlight and transfer it up to dark reaction. ATP together with NADPH2, called as assimilatory power and NADPH2 is called as reducing power.


It is also known as Blackmann’s reaction or thermochemical reaction. In this phase, the NADPH + H+ and ATP produced during light phase are used in the reduction or fixation of CO2 into carbohydrates. This reaction takes place in stroma of chloroplast 3.1.6.CALVIN CYCLE OR C3-CYCLE This method of CO2 fixation is described by Calvin, Benson and Bassham (1957). As first stable product of this reaction is phosphoglycericacid (PGA), which is a three carbon compound, this cycle is known to be as C3-cycle and the plants exhibit this cycle are called as C3 plants. Calvin has used unicellular algae Chlorella and Scenedesmus to study the C3 cycle. To identify intermediate compound he used radio tracer technique.


Summary of Calvin Cycle Carboxydismutase (i) 6RuDP + 6CO2 (RuDP-C) Kinase (ii) 12PGA + 12ATP Dehydrogenase (iii) 12 1.3-DPGA + 12ADP 12PGA 15 .3-DGPA + 12NADPH2 12 3-PGAld + 12NADP + 12H3PO4 12 1.

6-DP 5DHAP Phosphate 3F-6-P+3Pi Isomerase (x) 2 Ribose-5-P Epimerase (xi) 4 Xy-5-P Epimerase (xii) 2Ribose 2 Ribose – 5 – P Phosphatase (xiii) 6 Ribusole – 5 – P + 6 ATP 6 Ribusole-1.3 –PGAld Aldolase (viii)2E-4 –P + 2DHAP 2S-1.Isomerase (iv) 5PGAld Aldolase (v) 3PGA + 3DHAP Transketolase (vi) 1F-6-P 1Hexose Transketolase (vii) 2F-6-P + 2.7-DP + 2H2O 2S-7-P + 2H3P04 Transketolase (ix) 2S-7-P + 2PGAld 2Ribose-5-P+ 2Xy-5-P 2E-4-P + 2Xy-5-P Phosphatase 3F-1.6-DP 4 Ribulose – 5 – P 2Ribusole-5-P 16 .

temperature plays a very vital role. Bean. Chloroplast and mitochondria are also involved in this process. Main features of P.It is wasteful method and does not produce energy. Toxic H2O2 is formed during oxidation of the substrate 8.R. Rice. Photorespiration increases with the availability of O2 6.7PHOTORESPIRATION It has been observed that light affects respiration and the rate of respiration in light may be three to five times higher than the respiration in darkness. 9. It takes place in the presence of light. It is pronounced in C3 plants and negligible in C4 plants. 17 .Photorespiration takes place in peroxisomes.1. glycolate serves as substrate for photorespiration.3. etc.In normal respiration the respiratory substrate is sucrose which in photorespration glycolic acid (2 carbon compound) serve as a substrate. 4. It occurs in some plants like Beet. Such type of respiration is called photorespiration. 3. 7. its rate being very high in between 25-35˚C. Even up to 100%. are 1. 5.In photorespiration. It also depends upon the concentration of oxygen and increases with increasing oxygen concentration. End-products are CO2. 2.

18 .

1. etc. maize. M. Euphorbiaceae.Slack (1966) proposed an alternative pathway of CO2 fixation which is now known as Hatch and Slack pathway or C4-dicarboxylic acid pathway or C4-cycle or ß-carboxylation cycle. an alternate pathway to CO2 fixation in photosynthesis was discovered by Kortschak et al. and Nyctaginaceae.g. But in 1954. sugarcane. Occurrence : C4-cycle is found in the members of the family gramineae e.8HATCH AND SLACK CYCLE OR C4 – CYCLE Initially it was believed that CO2 fixation takes place only by Calvin cycle. who reported the formation of C4 dicarboxylic acid as primary product of photosynthesis in sugarcane. Characteristic Features of C4 plants : 19 .R.3. it is also found in the members of the family Cyperaceae. Azoaceae. in addition to Calvin cycle. *This cycle is known as C4 cycle because first stable product of this cycle is a four carbon compound known as oxaloacetic acid (OAA). Chenopodiaceae.D Hatch and C. Amaranthaceae.

Malic acid and aspartic acid is formed from pyruvic acid within mesophyll cells.B. (i) (ii) C4-cycle ( takes place in mesophyll cells) and C3-cycle ( takes place in bundle sheath cells) 20 . The leaves of C4 plants exhibit specific histological structure. Leaves of C4 plants contains two types (dimorphic) of chloroplast : (i) (ii) Mesophyll Chloroplast : It is smaller . 3. (ii) Phosphoenol pyruvic carboxylase (PEP-C): This enzyme is found in the chloroplast of mesophyll cells and it reduces atmospheric CO2 by C4 cycle.) of leaves of C4 plants is bounded by bundle sheath cells. This type of anatomical structure is known as Kranz anatomy. The vascular bundle (V. 2. Photorespiration does not take place in C4 plants or the rate of photorespiration is very low. lacking grana and possessing starch grains.1. Bundle sheath cells have different types of chloroplast. C4 plants have high photosynthetic rate (40-80 mg CO2 per hour) whereas the rate of photosynthesis in C3 plants is 10-15 mg CO2 per hour. The cells of bundle sheath are bounded by mesophyll cells. C4 plants are usually found in tropical region where temperature is between 3035ºC and light intensity is very high. 5. 7. (i) Ribulose diphosphate carboxylase or Rubisco : This enzyme is found within the chloroplast of bundle sheath cells. It catalyses the oxidative decarboxylation of malic acid to produce pyruvic acid and reduces CO2 to C3 cycle. grana is present there and starch grains are absent Bundle sheath Chloroplast: It is larger in size. There are two pathways of CO2 fixation in C4 plants. 6. 4.

Production in C4 plants is 2-3 times greater than C3 plants. First carboxylation reaction takes place in mesophyll chloroplast and second carboxylation takes place in bundle sheath cells in the following step wise reaction: Biological Significance of C4-cycle: 1. C4 plants possessing a very efficient enzyme system to utilize least amount of CO2. C4 plants can photosynthesize even in the presence of very low concentration of CO2.Mechanism of C4-cycle: There are two carboxylation reaction takes place in C4-cycle. This enzyme system is known as phosphoenol pyruvic carboxylase ( PEP-C). 2. 21 .

etc. These plants absorb CO2 during night and convert it into malic acid which is then stored in vacuoles. Monocot Families: Liliaceae. ------------------------------------. It was named as Crassulacean Acid Metabolism (CAM). 2. 4. write your answer in the space given. hence plants can even fix CO2 during short day conditions when very least concentration of CO2 is available. Compare your answer with that given at the end of the unit. 2. This CO2 is utilized by C3cycle. Asclepiadaceae.g.g. Since the cycle was first observed in the plants belonging to family Crassulaceae e. During day time (light) decarboxylation of malic acid takes place and CO2 is released. Kranz anatomy is found in-------------------------------------------plant 3. Opuntia) Azoaceae.9 CRASSULACEAN ACID METABOLISM OR CAM CYCLE It occurs mostly in succulent plants which grow under semi-arid conditions.3. Dicot Families: Crassulaceae e. PEP-C enzyme have high affinity with CO2 than RuDP-C enzyme. compositae.1.is essential for both photosynthesis and respiration 5. Chenopodium. (sedum. etc. 1. Sedum and Kalanchoe. Caryophyllaceae. Bryophyllum. Vitaceae. Euphoebiaceae. CHECK YOUR PROGRESS (1) Note: 1. This mode of CO2 fixation takes place during night (dark) because the stomata of leaves of these plants remain open only during night. The rate of photorespiration in C4 plants is very low (negligible) hence the rate of photosynthesis will be higher in these plants. An example of C4 plant is -----------------------------------------------------------------2. 22 . Similar metabolism has been reported in the plants belonging to following families: 1. CO2 acceptor compound in C3 plants is ----------------------------------------------4. Orchidaceae. convolvulaceae. P7oo is reaction centre of -----------------------------------------------------------------CO2 acceptor of C4 cycle is --------------------------------------------------------------- 3.

Thus. During day time decarboxylation of malic acid takes place and CO2 gas is released. Mechanism of CAM cycle CAM cycle is completed in following two parts: 1 Acidification and 2. As stomata opens during night.g. Acidification: Acidification takes place during following steps: (i) The stored carbohydrates are converted into phosphoenol pyruvic acid (PEP) through glycolysis. Malic acid formed during dark (night) is stored in large vacuoles.carboxylase PEP + HCO3Overall reaction is as follows : PEP .e. they fix atmospheric CO2 during night by CAM and fix internally borne CO2 by C3-cycle during day time. This CO2 is converted into sucrose and storage glucans (e. the CO2 diffuses freely into the leaf through open stomata at night. CO2 + H2O H2CO3 PEP. CO2 fixation takes place in chlorophyll containing cells of leaves and stem during night (dark) and malic acid synthesis takes place. (ii) The CO2 combine with PEP in the presence of phosphoenol–carboxylase (PEP-C) enzyme to produce oxaloacetic acid (OAA). 4. Pteridophytes: Polypodiaceae.3. The stomata remain closed during day (light) and open at night (dark). Characteristic Features of CAM plants 1. 3. Starch) by C3-cycle.C PEP + CO 2 + H2O OAA + H 3PO4 OAA + H3PO4 H+ + HCO3- 23 . CAM plants show diurnal cycle of organic acid formation i. 2. Deacidification 1.

takes place within living organisms and is generally catalyzed by enzymes. Deacidification: The decarboxylation of malic acid into pyruvicacid and CO2 in presence of light is called deacidification. Malate dehydrogenace OAA +NADPH2 Malic acid + NADP This malic acid. TC PA or C3 PGA Co2 CO2 liberated is fixed by C3 cycle on coming next night this starch is converted into PEP. In certain plants. During day (light) time malic acid stored in vacuoles is diffused out into the cytoplasm and become decarboxylated to produce pyruvic acid and CO2 in the presense of NADP malic enzymes(NADP-ME).10. One molecule of NADP is also reduced in this reaction. this reaction is catalysed by PEPcarboxykinase. 3. and is thus ready to accept atmospheric CO2. thus produced in dark as a result of acidification is stored in the vacuoles. Biosynthesis. The process is a vital part of metabolism.1.This reaction is facilitated in presence of reduced NADP+ ( =NADPH + H+) formed during glycolysis. The prerequisites for biosynthesis are: Precursor substances Energy (usually in the form of ATP) 24 . The oxaloacetic acid (OAA) may also be interconverted into aspartic acid 4. unlike chemosynthesis.(iii) The oxaloacetic acid (OAA) is not reduced into MALIC ACID in the presence of malic dehydrogenase enzymes. Biosynthesis of starch and sucrose Biosynthesis is a phenomenon wherein chemical compounds are produced from simpler reagents.

Other names sucrose in common use include UDPglucose-fructose glucosyltransferase. UDP-glucose: d-fructose-6-phosphate glucosyltransferase. vitamins. The systematic name of this enzyme class is NDP-glucose:Dfructose 2-alpha-D-glucosyltransferase.13) is an enzyme that catalyzes the chemical reaction NDP-glucose + D-fructose NDP + sucrose Thus. Biosynthesis of sucrose The enzymes that catalyze sucrose biosynthesis and cleavage in higher plants were first reported by Cardini et al. UDP-glucose: d-fructose-2-glucosyltransferase. Starch.3. Important and commonly-known products of biosynthesis include proteins. and antibiotics. The knowledge of sucrose metabolism in unicellular organisms is limited. Enzyme: Sucrose synthase In enzymology.14).13) were isolated and partially purified from wheat germ.00). sucrose-6-phosphate phos phohydrolase. specifically the hexosyltransferases. NADPH. a sucrose synthase (EC 2. sucrose-uridine diphosphate glucosyltransferase. and others). EC 2.4. usually enzymes Reduction equivalents (in the form of NADH. 25 .4. and sucrose synthase (SS.Often required components include: Catalysts. the two substrates of this enzyme are NDP-glucose and D-fructose. it may be possible. to harness this process for the production of biodegradable plastics. This enzyme belongs to the family of glycosyltransferases. and uridine diphosphoglucose-fructose glucosyltransferase. but all components of living beings are a result of this process eg.2. sucrose-UDP glucosyltransferase.4. whereas its two products are NDP and sucrose. its specific phosphatase (SPP. In the near future.1. EC 2. sucrose and lipids. EC 3. with the help of biotechnology.1. and Leloir and Cardini in 1955.1. Sucrose-phosphate synthase (SPS. This enzyme participates in starch and sucrose metabolism. synthetase.

2-biphosphate is hydrolysed to fructose-6-phosphate.An outline of sucrose biosynthesis is as follows: Fructose 1.6-hosphate. Glucose-6-phosphate ultimately gives rise to UDP-D-glucose. This ultimately produces sucrose. This splits into fructose and glucose. 26 .

The formation of the starch granule can be viewed as a simple model for the formation of ordered three-dimensional polysaccharide structures in plants. each -1000 residues long. Starch is synthesized in leaves during the day from photosynthetically fixed carbon and is mobilized at night. this issue). 1995. fruits. Amylose consists of predominantly linear chains of a(l4)-linked glucose residues. Amylose is usually branched at a low leve1 (approximately one branch per 1000 residues) by a (1-6) linkages and makes up -30% of starch. It is also synthesized transiently in other organs. degree of polymerization. It consists of different glucose polymers arranged into a threedimensional. the universality of its distribution among different plant species. However. such as meristems and root cap cells. Understanding the biochemical basis for the assembly of the granule could provide a conceptual basis for understanding other higher order biosynthetic systems such as cellulose biosynthesis (see Delmer and Amor. it may crystallize and shrink (retrogradation) after heating (Shewmaker and Stalker. which in storage organs committed primarily to starch production are called amyloplasts. For example. THE STRUCTURE OF STARCH AND THE STARCH GRANULE Starch can be chemically fractionated into two types of glucans polymer: amylose and amylopectin. semicrystalline structure-the starch granule. 1992). amylose forms hydrogen bonds between molecules. Starch is synthesized in plastids. Once extracted from plants and in solution. The biosynthesis of starch involves not only the production of the composite glucans but also their arrangement into an organized form within the starch granule. and storage roots. including seeds. tubers. depending on the concentration. resulting in rigid gels. one emerging concept is that structure within the granule itself may determine or influence the way in which starch polymers are synthesized. and temperature.Starch Biosynthesis Starch is the most significant form of carbon reserve in plants in terms of the amount made. but its major site of accumulation is in storage organs. which consists of highly branched 27 . and its commercial importance. Amylopectin.

glucans chains, makes up -70% of starch. Chains of roughly 20 a(l4)-linked glucose residues are joined by a(1-6) linkages to other branches. The branches themselves form an organized structure (Figure 1A). Some are not substituted on the six positions and are called A chains. These chains are a(1-6) linked to inner branches (B chains), which may be branched at one or severa1 points. A single chain per amylopectin molecule has a free reducing end (the C chain). The branches are not randomly arranged but are clustered at 7- to 10-nm intervals (Figure 1). An average amylopectin molecule is 200 to 400 nm long (20 to 40 clusters) and -15 nm wide (for review, see Kainuma, 1988; Smith and Martin, 1993). After extraction, amylopectin has more limited hydrogen bonding than amylase in solution and is more stable, remaining fluid and giving high viscosity and elasticity to pastes and thickeners. Some starch, most notably that from potato tuber, is also phosphorylated.



Figure 1. Amylopectin Structure, Starch Granule Form, and Starch Biosynthesis. (A) Diagrammatic representation of an amylopectin molecule. a(M)-Linked glucans are attached by a(1-6) linkages to form a highly branched structure. Short glucan chains (A chains) are unbranched but linked to multiple branched B chains. There is a single reducing end to the C chain glucan. The branches are arranged in clusters 4 0 nm long, with a few longer chains linking more highly branched areas. (B) Diagrammatic representation of a starch granule from storage tissue showing alternating semicrystalline and amorphous growth rings. The semicrystalline regions are thought to consist of alternating crystalline and amorphous lamellae. (C) Steps of starch biosynthesis. ADPGPPase catalyzes the formation of ADPglucose and inorganic pyrophosphate from glucose-I-phosphate and ATP (step 1). Starch synthases (SS) add glucose units from ADPglucose to the nonreducing end of a growing a(l+linked glucan chain by an a(l-4) linkage and release ADP (step 2). Starch-branching enzymes (SBE) cut an a(l-4)-linked glucan chain and form an a(1-6) linkage between the reducing end of the cut chain and the C6 of another glucose residue in an a(lr()-linked chain, thus creating a branch (step 3).

THE BIOCHEMISTRY OF STARCH BIOSYNTHESIS The biosynthetic steps required for starch biosynthesis are relatively simple, involving three committed enzymes: ADPglucose pyrophosphorylase (ADPGPPase; EC, starch synthase (SS; EC, and starch branching enzyme (SBE; EC; Figure 1C). 60th amylose and amylopectin are synthesized from ADPglucose, which is synthesized from glucose-1-phosphate and ATP in a reaction that is catalyzed by ADPGPPase and that liberates pyrophosphate. This enzyme is active within the plastid, which means that its substrates, glucose-1-phosphate and ATP, must also be present in the plastid. In chloroplasts, ATP may be derived from photosynthesis, but in nonphotosynthetic plastids, it must be specifically imported from the cytosol, probably by an ADP/ATP translocator (Ngernprasirtsiri et al., 1989; Schünemann et al., 1993). The 30

causing the release of ADP. as discussed previously. VARIABLE PARAMETERS IN STARCH BIOSYNTHESIS The relative simplicity of the starch biosynthetic pathway does not explain the enormous variability in starch composition among different plant species. In all species that have been investigated. Branches are not created randomly. which hydrolyzes an a(l-4) linkage within a chain and then catalyzes the formation of an a(1-6) linkage between the reducing end of the “cut” glucan chain and another glucose residue. In nonphotosynthetic tissues. 1987). probably one from the hydrolyzed chain. The removal of this plastidial pyrophosphate effectively displaces the equilibrium of the ADPGPPase reaction in favor of ADPglucose synthesis (weiner et al. We are only just beginning to understand how these layers of complexity are determined. and tissues. 31 . it may be imported directly from the cytosol (Tyson and ap Rees. but already it is clear that central to their organization is diversification in the activities of the participating enzymes and modulation of the extent of their activities. The a(1-6) branches in starch polymers are made by SBE. In the next step of starch synthesis. 1988) or synthesized in the plastid from glucose. SS catalyzes the synthesis of an a(1-4) linkage between the nonreducing end of a preexisting glucan chain and the glucosyl moiety of ADPglucose.glucose-1-phosphate can be supplied by the reductive pentose phosphate pathway in chloroplasts via phosphoglucoisomerase and phosphoglucomutase (Smith and Martin.6-phosphate via the action of a plastidial phosphoglucomutase (Hill and Smith. 1993). 1991). SSs can use both amylase and amylopectin as substrates in vitro. SBEs show some specificity for the length of the a(1-4)glucan chain that they will use as a substrate.. Nor does it explain the complexity of starch in terms of its component glucan chains and their organized arrangement in starch granules. The pyrophosphate produced by ADPGPPase is removed by inorganic alkaline pyrophosphatase. a structure that requires a minimum glucan chain length. but show an average periodicity of 20 glucan residues. Part of this selectivity may reside in the fact that these enzymes cleave only those glucan chains that are in a stable double helical conformation. which is probably confined to plastids in both photosynthetic and nonphotosynthetic tissues. varieties.

1993. Mu et al. Consequently. 1993.. These assignments allow not only the comparison of the roles of particular isoforms from different species but also a clearer view of how starch biosynthetic gene expression is controlled and the contribution this control makes to the overall regulation of starch biosynthesis. and several reports indicate that the potential for allosteric regulation of ADPGPPase from some storage organs is relatively insignificant in comparison with that in leaves (Hylton and Smith. The existence of isoforms clearly provides flexibility for specialization and control in starch biosynthesis.. for example. their time of expression during starch granule formation. because protein degradation is a common feature of purification of SSs and SBEs (see. 1991... Changes in the levels of 3-PGA and Pi in leaves are modulated primarily by the rate of photosynthetic carbon fixation. In leaves. Kleczkowski et al. which has begun to allow the assignment of isoforms to particular families with related primary structures and apparently related functions. The understanding of the roles of different isoforms is therefore being greatly facilitated by molecular analysis. There is strong evidence that changes in rate are achieved through allosteric regulation of ADPGPPase by the activator 3-phosphoglycerate (3-PGA) and the inhibitor inorganic phosphate (Pi). 1994) and could indeed be of significance in vivo. 1991). One problem in characterizing isoforms for each step has been that their identification has been based predominantly on activities in biochemically fractionated extracts. Weber et al. However. it is unlikely that these are all the products of different genes. 1992. Baba et al. Starch biosynthesis in many storage organs does not have an obvious requirement for short-term metabolite-mediated regulation. It is possible that the contribution of metabolic regulation to starch biosynthesis may vary across the plant. thus giving rise to significant modulation of ADPGPPase activity (Preiss. Blennow and Johansson. control of 32 . These may differ in their products.there are isoforms for each of the committed steps of starch biosynthesis. Some modulation of starch biosynthesis can also be achieved through metabolic control of flux through the pathway. starch synthesis occurs at higher rates when carbon assimilation is high relative to the demand for carbon export and at lower rates when assimilation is low relative to demand from the rest of the plant. This approach has allowed many proteins with either SS or SBE activity to be characterized from different species. their kinetic properties. 1995). and the organs in which they are active.

More recent analysis of SSs has shown that the biochemical distinctions are not absolute. even in tissues in which allosteric regulation is not important.starch biosynthesis may also involve modulation of the extent of allosteric regulation of ADPGPPase. however. However. SSs found in the soluble phase may also be bound to the granule (Denyer et al.empirically in different tissues and under different conditions. it is difficult to assess the importance of such "noncatalytic" properties without undertaking lengthy in vivo assays using mutagenesis and plant transformation. 1993. the activity of ADPGPPase may exercise significant control in starch biosynthesis. For example. the presence of particular noncatalytic characteristics in different forms of each biosynthetic enzyme potentially represents another way in which the pathway can be diversified to give more complexity in glucan products and their organization and more variation among different plant species. "noncatalytic" characteristics are probably still of functional significance because they could dictate how active a particular isoform may be when bound to the granule. which can have a profound effect on the products made within the starch granule. this needs to be tested. However. 1995). which may not reflect precisely the conditions within the granule. most assays of starch biosynthetic enzymes have been made on soluble or solubilized extracts. This is most clearly seen for SSs. whereas soluble SSs would synthesize amylopectin. This is likely due 33 . Therefore. a functional distinction was predicted. Of course. indicating that its synthesis is somewhat delayed compared with that of amylopectin (Shannon and Garwood. 1984). that granule-bound SSs (GBSSs) would synthesize amylose. which are located both bound to the starch granule and in the soluble phase of the amyloplast. To date. The different proteins involved in starch biosynthesis may also vary in their physical characteristics. Following biochemical analysis of waxy (wx) mutants from several species. Starch biosynthesis varies both quantitatively and qualitatively during the course of storage organ formation. namely. Some isoforms of the biosynthetic enzymes may be active early in starch granule formation and others active later. amylose content normally increases as a proportion of total starch during storage organ development..

. Whereas some species show significant differences in expression of starch biosynthetic genes at different developmental stages (Dry et al. the effective activity of the biosynthetic enzymes at different developmental stages in vivo is very difficult to assess. 1992. others show few differences (Kossmann et al.partly to the timing of production of the amylose-specific GBSS.... Hauxwell et al. 1990). 1991. This apparent lack of developmental change could be an artifact of the way gene expression is assayed. Nakamura and Yuki. Starch Biosynthesis 975 1993). 1978. 1989. This means that the turnover of these enzymes is relatively low. waves of gene expression have been reported across developing seed from more advanced to less advanced cells (Shannon and Garwood. the use of molecular biology and genetics has complemented the biochemical analysis of this system to allow a greater appreciation of the control of each biosynthetic step and its contribution to the overall process. Photosynthesis: Physiological and Ecological Considerations 34 .. For example. However. which is synthesized later than some other SSs (Nelson et al. In fact. 1984). it is unclear to what extent the trapped enzymes are active within the granule. As the starch granule grows. Perez-Grau and Goldberg. A final factor complicates the potential significance of developmental modulation in the control of starch biosynthesis. There may also be developmental gradients in starch biosynthesis within a storage organ (Shannon and Garwood.. Mizuno et al. Burton et al. Dry et al. Although our understanding is far from complete. 1995). thus. it may be that developmental regulation of isoform gene expression is more important than is appreciated at present. 3.2. storage organs with strong interna1 developmental gradients tend to "flatten out" developmental differences in assays based on total extracts. 1995). 1984. 1993.. 1992).. 1992. The plant can thus use severa1 different strategies to refine starch biosynthesis and to build the organized form of the granule. many of the biosynthetic enzymes become trapped within it (Denyer et al.

Units can be expressed in terms of energy. Concepts and units for the quantification of light Amount of light. The first two parameters. where "moles" refers to the number of photons (1 mol of light = 6. Light can also be thought of as a stream of particles. amount and direction. The energy of a photon is related to its wavelength as follows: 35 . and the amount of energy that falls on a flat sensor of known area per unit time is quantified as irradiance. In this case. Quanta and energy units can be interconverted. Working with Light Three light properties are especially important when working with light: amount. direction. such as watts per square meter (W m-2).From physiological and ecological view point. a planar light sensor is the most appropriate. photons. Is the plant part flat or cylindrical? For a flat leaf. when considered as a wave. This measure is called photon irradiance. carbondioxide concentration and temperature have following considerations to be studied. Avogadro's number). Time (seconds) is contained within the term watt: 1 W = 1 joule (J)s-1. are important with respect to the geometry of the part of the plant that intercepts the light. units can be expressed in moles per square meter per second (mol m–2 s–1). provided that the wavelength of the light is known. light has a wavelength and a frequency. in order to understand how photosynthesis responds to environmental factors like light.02 × 1023 photons. and spectral quality. The energy of a photon depends on its frequency. as expressed by Planck's law. or quanta.

A photon of 400 nm light contains 4.48 × 10-19 J. When light deviates from perpendicular.63 × 10–34 J s). Thus. the higher the wavelength of a photon.988 × 10-16. usually expressed in nm (1 nm = 10–9 m). We can solve for the hλ part of the equation. the 800 nm photon contains 2. and it decreases as light becomes more oblique—similar to the situation with a typical leaf. and λ is the wavelength of light. as indicated by the larger denominator in the equation. On the other hand.where c is the speed of light (3 × 108 m s–1). 36 . Turning our attention to the direction of light. Sensors that correct for the angle of incidence of light are said to be cosine corrected. which is in the blue region of the spectrum. From this equation we can see that a photon at 400 nm. has twice the energy of a photon at 800 nm.97 × 10–19 J. irradiance is maximal when light strikes a surface directly from above. irradiance is proportional to the cosine of the angle at which the light rays hit the sensor. and we obtain 1. from the infrared region of the spectrum. Direction of light. h is Planck's constant (6. the lower its energy. and write this equation as: where λ is expressed in nanometers. Stated differently. light can strike a flat surface directly from above or it can strike the surface obliquely.

soil.. whole plants. chloroplasts). complex shoots. In addition. Equivalent amounts of collimated light strike a flat irradiance-type sensor (A) and a spherical sensor (B) that measure fluence rate. direct light from the sun plus the light that is reflected upward from sand. A and B will give the same light readings.g. the type of measurement is called fluence rate (Rupert and Letarjet 1978). or snow). the flat irradiance sensor (C) will measure an amount equivalent to the irradiance in A multiplied by cosine of the angle α in C. In contrast.g. (After Björn and Vogelmann 1994. and the measured amount of light can be expressed in watts per square meter (W m–2) or moles per square meter 37 . In these situations it makes more sense to measure light with a spherical sensor that measures light omnidirectionally (from all directions). Irradiance and fluence rate. the spherical sensor (D) will measure the same quantity as in B.Figure. When the amount of light is measured by this omnidirectional measurement.. With collimated light.) There are many examples in nature in which the light-intercepting object is not flat (e. in some situations light can come from many directions simultaneously (e. When the light direction is changed 45°.

Moreover. Flat. or SI units) "density" can mean area or volume. it has been suggested that the term "density" be discontinued (Holmes et al. it is often given the special term photosynthetic photon flux density (PPFD). area is contained within the term flux. It is clear from the units whether light is being measured as energy (W) or as photons (mol).25 times the fluence rate. PPFD has in some cases been shortened to PPF. When light is completely diffuse. values for fluence rate versus irradiance can be quite different from one another. but it is not clear whether this abbreviation represents an irradiancetype or a spherical measurement. In research on photosynthesis. cosine-corrected sensors are ideally suited to measure the amount of light that strikes the surface of a leaf. The corresponding value in energy units is about 400 W m–2. PAR irradiance and fluence rate are both about 2000 µmol m–2 s–1. Depending on whether the light is collimated (rays are parallel) or diffuse (rays travel in random directions). 400 to 700 nm) may also be expressed on the basis of energy (W m–2) or quanta (mol m–2 s–1) (McCree 1981). How much light is there on a sunny day and what is the relationship between PAR irradiance and PAR fluence rate? Under direct sunlight. It is important to note that PAR is an irradiance-type measurement. However. 38 . a spherical sensor is equally sensitive to light from all directions. The flat sensor measurement of photosynthetically active radiation (PAR. 1985) because within the International System of Units (Système Internationale d'Unités. though higher values can be measured at high altitudes. irradiance is only 0. such as when studying a chloroplast suspension or a branch from a tree. when choosing how to quantify light. In contrast to a flat sensor.per second (mol m–2 s–1). spherical sensors are more appropriate in other situations. see Björn and Vogelmann 1994 and Kirk 1994). it is important to match sensor geometry and spectral response with that of the plant. In summary. They are equivalent only under special conditions (for a detailed discussion. when PAR is expressed on a quantum basis.

0. 0. and the ratio of the two is called the Bowen ratio (Campbell 1977): This concept was developed by Ira S. by sensible (or perceptible) heat loss.8 for temperate forests and grasslands.2 for tropical rain forests and 0. the Bowen ratio is about 10 for deserts. On the other hand. 2-6 for semi-arid regions. the heat loss from the soil. Thus. because water supply is limited. One can calculate the evapotranspiration rate for an entire canopy using measurements of the Bowen ratio. 39 . a leaf with an effective thickness of water of 300 µm would warm up by 100°C every minute if all available solar energy were absorbed and no heat was lost. net incident radiation.Heat Dissipation from Leaves: The Bowen Ratio The heat load on a leaf exposed to full sunlight is very high. Plants with very high Bowen ratios conserve water but have to endure very high leaf temperatures in order to maintain a sufficient temperature gradient between the leaf and the air. so the Bowen ratio is low. and the gradients in temperature and water vapor concentration above the canopy (Ibanez and Castellvi 2000).1 for tropical oceans (Nobel 1999) In well-watered crops. However. is high. this enormous heat load is dissipated by the emission of long-wave radiation. and the Bowen ratio is infinite. stomata closure prevents evaporative cooling. an American astrophysicist. the Bowen ratio tends to be high. In fact. and by evaporative (or latent) heat loss.4 to 0. in some cacti. all the heat is dissipated by sensible heat loss. When the evaporation rate is low. Bowen (1898–1978). Slow growth is usually correlated with these adaptations. transpiration (and hence water evaporation from the leaf. Sensible heat loss and evaporative heat loss are the most important processes in the regulation of leaf temperature.

Climate is a major factor influencing the natural distributions of C3 plants and C4 grasses. so the important factor influencing photosynthetic pathway distribution becomes the temperature during the growing period. Here. and not found in trees (with a single Hawaiian tree exception. Based on many systematic surveys of the natural vegetation across the globe that have been accumulating over time. Below we construct a global map that describes the general abundances of C3 and C4 taxa on different continents. less common in herbs and shrubs.The Geographic Distributions of C3 and C4 Plants Among the 15. it is most common in grasses and sedges. C4 taxa are found in warm to temperate environments and are uncommon in cool to cold climates. Euphorbia forbesii). a clear picture is emerging. Clearly plants will not grow in the absence of water. we talk about the two most important climate parameters influencing plant growth: water and temperature. 40 .000+ species with C4 photosynthesis.

the two dominant C4-photosynthesis subtypes do not share identical distributions. because dense tropical forests tend to shade out C4 grasses. Cerling. triangular presentation of the different gradients influencing the abundances of different C4 photosynthesis subtypes. Courtesy of Ehleringer. This reflects the role of disturbances. Courtesy of Ehleringer. wetter edge of the Great 41 . such as grazing and fires. and Dearing (2005). The C4 NADP-me grasses tend to occur in drier regions. Human activities and disturbances by animals will influence the distribution of C3 and C4 taxa within savannas and grasslands. The C4 taxa are most common away from the tropics in the savanna and steppe regions. their abundances tends to diminish south of the desert zones (generally 30–40° latitude). Notice that C4 taxa are not very common in tropical regions (0–20° latitude). Across the grasslands and savannas. Cerling. and Dearing (2005). on the importance of trees on the landscape. A three-axis. such as the shortgrass prairie of the Great Plains.Figure. A map of the geographical abundances of C3 and C4 grasses in the savannas and grasslands of the world. On the eastern. Figure.

We measure atmospheric CO2 in units of ppm or parts per million. Projected Future Increases in Atmospheric CO2 Human use of fossil fuels (coal. agricultural practices result in C4 plants growing outside of the distributions shown above. A high precision record of the atmospheric carbon dioxide levels measured on Manua Loa. Courtesy of NOAA. Earth System Research Laboratory: Note the cyclic nature of the atmospheric CO2 data. and millet in both tropical and temperate regions. sugarcane. and natural gas) continues to increase as growing human populations demand more energy for transportation. Hawaii. This annual pattern reflects changes in the balance of photosynthesis (decreases atmospheric CO2) and respiration (increases atmospheric CO2) at a location over the course of the year. heating. the shortgrass prairie is replaced by tallgrass prairie. oil.Plains. Today. in which one oscillation cycle is exactly one year. dominated by C4 NAD-me grasses. This results from the extensive planting of corn. The rate of atmospheric increase in CO2 is about 3 ppm per year Figure. and manufacturing. Atmospheric CO2 tends to decrease in the spring and 42 .

One surprising fact is that the observed rate of atmospheric CO2 increase is actually less than the observed rate of atmospheric CO2 increase. Global warming and changes in climate are anticipated effects of a rapidly increasing CO2 levels.e. it is doubtful that plant growth can be sustained in a linear. 43 . These FACE research facilities give scientists an opportunity to understand how different plant biochemical. proportional fashion as atmospheric CO2 levels continue to increase.summer. The U. The FACE studies are designed to address the question of how ecosystems will respond to future atmospheric CO2 environments and whether the growth response level off at some future CO2 level. European countries. of course. pipes inject CO2 into the interior of a ringed area containing a complete ecosystem as shown below. Economists have good estimates of the rate of CO2 emission globally.S. and growth processes within the ecosystem will respond as a result of long-term exposure to elevated CO2 levels. This is because plants on land and algae in the ocean are currently able to take up about one-half of fossil fuel emissions through enhanced photosynthesis.. In 2007 atmospheric CO2 reached an average value of 384 ppm and is expected to reach 400 ppm before 2015. physiological. mineral nutrients are required as well). Scientists study how plants and ecosystems respond to elevated CO2 using an experimental field approach called a Free Air CO2 Enrichment (FACE) Experiment. China. based on two plausible scenarios. Below are estimates of the ranges. These are.. atmospheric CO2 tends to increase in the fall and winter when respiration rates exceed photosynthesis. In a FACE experiment. Since biomass production involves so much more than simply increased photosynthesis (i. when photosynthesis rates within an ecosystem exceed respiration rates. Japan. and India are the largest sources of fossil fuel emissions. In contrast. Just how much the atmospheric CO2 will increase is unknown. but two of the many reasons why scientists and others are concerned about the consequences of elevated atmospheric CO2.

However. muscle. 44 . The rate of photorespiration in a leaf increases because RuBP carboxylase is more likely to react with O2 instead of CO2 as temperature increases and/or CO2 decreases. The vast majority of plants do not become fossils under these conditions. Below we provide a 3-D graphic of how the ratio of photorespiration-to-photosynthesis increases as temperature increases and/or CO2 decreases. it is possible to choose from among. bone collagen. but instead are decomposed by microbial activities leading to diminished numbers of fossils that we can use to reconstruct the expansion of C4 plants over time. Hair provides a sequential record of the animal’s diet during the time of hair production. there are good proxies. titled “business as usual” atmospheric CO2 levels are projected to reach 700 ppm by the end of this century. because individual plants are not well recorded in the fossil record in many locations. an aggressive global effort to curb CO2 emission might result in an atmospheric CO2 stabilization of 550 ppm. and tooth enamel for isotope analyses in order to determine the proportions of C3 and C4 food sources in their diets. The result of an increase in photorespiration is a decrease in photosynthetic quantum efficiency. aquatic environment. Reconstruction of the expansion of C4 plants over the past 10–15 million years can be a challenge. For modern animals. This leads to environmental conditions where C4 plants are favored over C3 and vice versa.In one scenario. On the other hand. Reconstruction of the Expansion of C4 Taxa C4 photosynthesis is favored over C3 photosynthesis under conditions of high temperature and/or low atmospheric CO2. Most plant fossils are thought to have formed as a result of submersion into an anaerobic. hair. Carbon isotope ratios (δ13C) of animal tissues reflect the foods that they ate.

As teeth are preserved for millions of years. Shown below are the time series analyses of carbon in bog in tropical Africa. about the only remaining tissue that remains and preserves the C3/C4 dietary signals is the enamel in the animal’s teeth. C4 photosynthesis is predicted not to occur on Earth until the atmospheric CO2 level falls below a critical threshold value. For many herbivores. the vegetation surrounding these bogs switched from being dominated by C4 plants to being dominated by C3 plants. There is also extensive evidence presently to show that fluctuations in the atmospheric CO2 concentration values between glacial (180 ppm) and interglacial (280 ppm) periods were large to influence the local abundances of C3/C4 plants. δ13 of tooth enamel can be used to reconstruct the abundances of C3 and C4 plants eaten by mammalian grazers over extended time periods. C4 plants were not an important part of ecosystems. between 6 and 9 million years ago.The δ1313C of animal teeth faithfully record the carbon isotope ratios of food sources during the period of tooth development. Then. such as cows and horses. C4 plants became important parts of many ecosystems. This is the isotope enrichment associated with CO2 condensing to form carbonate. The carbon in tooth enamel is actually carbonate that has condensed from CO2 that was produced as a byproduct of metabolizing food. Until 8 million years ago. After an animal has died and decomposition has occurred. particularly at tropical and semi-tropical latitudes. Thure Cerling and his colleagues at the University of Utah have applied the “tooth enamel” tool to reconstruct the historical abundances of C3/C4 plants.0000 to 12.000 years ago.Note that there is an “ε” offset of 14. tooth growth is continuous and provides a longer-term dietary record than for animals with a fixed tooth growth period (such as humans). At about 10. These results are consistent with model predictions above. This shift is evident by the dramatic change is carbon 45 .1‰. Now we have a tool—the carbon isotope ratio of animal tooth enamel will record the abundance of C3 versus C4 food sources. Even then it should appear first in those locations with the warmest temperatures during the growing season.

2. Q1. kranz anatomy Q2. especially at the time that the glacial period ended and we entered our current inter-glacial period. CHECK YOUR PROGRESS (2) Note: 1. The C3 photosynthetic pathway dominated for much of Earth’s history. since anthropogenic burning of fossil fuels is rapidly increasing in atmospheric CO2 to levels far exceeding those observed on Earth over the past 1 million years. Distinguish the process of CO2 reduction between C3.isotope ratios of materials feeding into the bog. Q3. write your answer in the space given. Compare your answer with that given at the end of the unit. Draw diagram of chloroplast. And it is only relatively recently in Earth’s (6–8 million years ago) history that we have seen an expansion of C4dominated ecosystems. What the future holds is unclear. Together these historical pieces of information paint an interesting history. Write short notes on:(a) Photorespiration or C2-cycle (b) calvin cycle (c) photolysis of water (d) Non – cyclic photophosphorylation (e). C4 and CAM plants. Kranz type of anatomy 46 .

cytochrome 5. and leads to generation of precursor metabolites reducing power. PEP Check your progress. An out line of cycle & main features. 3. The energy required for biological activities is obtained from organic compounds available in food. 1. RuDP 4.reduction process.Key I Answers. Maize 2. + NADPH+H +ATP.Key II Your answer must include 1. etc get successively oxidized to produce CO2. C4 plants 3.Check your progress. all living organisms respire to produce energy needed to perform all vital activities. An out line of there pathways and differentiation among them .” 47 . To recall. H2O and energy. 2. The process of respiration is basically an oxidation. fats.3RESPIRATION__________________________________________________ INTRODUCTION: In this unit you will learn about the process of respiration in plants. where electrons are withdrawn from substrate (glucose) are accepted by various components of etc( electron transport chain] and reducing powers. Defination : “ Respiration is a process by which organic food materials such as sugar. Plants synthesize their own food through photosynthesis. PSI 6.

ADP + H3PO4 ATP(ADP˜P) Energy stored in ATP is utilized for carying out different cellular and biological activites because of this. 48 6CO2 + 6H2O + 38ATP . • b) O2 utilized in the process comes through stomata &CO2 is The sites of respiration are cytoplasms and mitochondria. The energy released is stored in pyrophosphate bonds of ATP.C6H12O6 + 6O2 6CO2 + 6H2O + 673Kcal energy The overall reaction of cellular repiration is given as C6H12O6 + 6O2 + 38Adp +38iP Objective: The main aim of this unit is to develop an understanding of the process of respiration. anaerobic respiration & fermentation. compounds are broken down inside the cells by oxidation process. After learning this unit you will be able to • • Differentiate between various types of (respiration.1. known as cellular respiration. fermentation) fueling reaction. AN OVERVIEW OF PLANT RESPIRATION. Understand the significance of respiration in a) Generation of precursors b) Generation of reducing power c) Generation of ATP • • • Realize the role and significance of various enzymes involved in the process. a) You must bear a clear understanding in mind that both photosynthesis and respiration involves gaseous exchange but light reaction of photosynthesis requires sunlight whereas respiration occurs all the time. energy is called energy currency of the cell. Understand the existence of alternative oxidation pathways The applications of fermentation and the basic difference between the process of aerobic respiration . The organic released through the same.3. 3.

it is completed in 3 steps: a) Glycolysis / EMP pathway b) Oxidation of pyruvic acid c) ETC & oxidative phosphorylation 3. fructose. If I say that glycolysis is a fermentive pathway would you agree? 49 6CO2 + 6H2O + 38ATP . The ratio of volume of CO2 released to the volume of O2 absorbed during respiration is called respiratory ratio or R.2GLYCOLYSIS/ EMP PATHWAY In Greek language the word glucose means sugar and lysis means dissolution. protein.Q. a particular enzyme is required which works in a sequential manner one after the another. = Volume of O2 absorbed To develop a clear understanding of the process let us understand the mechanism of respiration MECHANISM OF RESPIRATION Cellular respiration is a complicated process which is completed in many steps. Volume of CO2 released R. fats.3.Q. for every step.c) The overall reaction is as follows: C6H12O6 + 6O2 + 38Adp +38iP The main features of respiration in plants are: • • • • • • • • Oxidation of organic compounds occurs in under aerobic conditions Complete oxidation occurs End products are CO2 & H2O Higher amount of (673 Kcal )energy is liberated out Process occurs in cytoplasm and mitochondria Chlorophyll pigment is not essential Various respiratory substance are: glucose. etc.

Reasons to support my statement are: a) It does not involves O2 intake b) ATP generated is through substrate level phosphorylation. This process was discovered by three German scientists Embden. Thus the process of sequential oxidation of glucose into pyruvic acid is known as glycolysis. 50 . All the reactions of glycolysis take place in the cytoplasm and through the glycolysis glucose is oxidized into pyruvic acid in presence of many enzymes present in the cytoplasm. meyerhof and Parnas. c) Organic compound donates electrons and organic compound accepts it. On their name the pathway is also called EMP pathway.

3-DPGA 3-PGA Pyruvic Acid No.3.7.P~P + P adenosine ~P + P adenosine + P 4° = .3Kcal 4° = .3 – DPGA PEPA 1.P ~ P + H2O Adenosine ~ P + H2O adenosine .P ~ P ~ P + H2O Adenosine . (One molecule of NADH2 gives three molecules of ATP by ETC) Total production of ATP in glycolysis cycle Reaction number (vii) (viii) (xi) 1.Energy production during glycolysis: During glycolysis process two molecules of ATP are utilized to convert glucose into glucose-6-PO4 & fructose -1.3 –DPG-Ald 1.4Kcal High energy compounds other than ATP 51 . of ATP molecule produced 2NADH2(2*3) = 6ATP 2ATP 2ATP = 2ATP = 2 ATP 10ATP As 2 molecules of ATP are utilized during glycolysis. thus net gain of ATP molecules during this process is 8 molecules of ATP 10 ATP – 2 ATP Net gain of ATP = 8 ATP SIGNIFICANCE OF GLYCOLYSIS: a) Generate ATP b) Precursor metabolic generation c) Generates reducing power Main enzymes are: 1) phosphofructokinase 2) pyruvate kinase 3) pyruvate enol carboxylase General patter of metabolism leading to synchronization in Ecoli cells Role of ATP : • • • Adenosine .8Kcal 4° = .7. 6 diphosphate where as 4 molecules of ATP and 2 molecules of NADH2 are produced during following steps.

A.C.(1942) and H. Pyruvic Decarboxylase CH3COOH CH3CHO + O2 2) In presence of alcohol dehydrogenase enzyme acetaldehyde reacts with NADH2 to produce ethyl alcohol and NAD.OH + 2NAD Ethylalcohol . A.al. Pyruvic Decarboxylase 52 CH3. H2O and ATP. Before entering Kreb’s cycle pyruvic acid gets decarboxylated to produce acetyl-CoA which enters the Kreb’s cycle and oxidize to produce CO2 .dehydrogenase 2CH3.Kreb’s (1943) in the presence of O2 oxidation of pyruvic acid takes place through Kreb’s cycle or T.CH2.of: Protein(ribosome function) Phospholipids Peptidoglycan layer of bacterial wall lipopolysaccarid layer of bacterial wall Fatty acids The fat of pyruvic acid produced during glycolysis depends on whether oxygen is available or not A) In case of anaerobic condition it is used as hydrogen acceptor for the two molecules of NADH generated during glycolysis and is converted into lactic acid. Alcoholic fermentation of pyruvic acid in plants: in yeast cells anaerobic oxidation of pyruvic acid takes place as follows: 1) Decarboxylation of pyruvic acid in presence of pyruvic decarboxylase enxyme to produced acetaldehyde.CHO + 2NADH2 Acetaaldehyde In animal cells lactic acid is formed B) Aerobic oxidation of pyruvic acid according to Wood et.Compound GTP CTP UTP Dcoxythymidine~ P~P~P dTTTP Acyl~SCoA OXIDATION OF PYRUVIC ACID cause action in Priosyn.A cycle.

Pyruvic acid + Glutamic acid 3.T. CYCLE/KREB’S CYCLE: Acetyl-CoA +NADH2 C) Fate of pyruvic acid to alanine during amino acid synthesis pyruvic acid react with Alanine +α-Keto glutaric acid This cycle was described for the first time by H.3. cycle because it produces tricarboxylic acids the process completes in mitochondrial crests.A.3.C. It is also known as T.C.Pyruvic acid + Coenzyme A + NAD glutamic acid alanine.A.Kreb’s in 1943.A. 53 .

Overall reaction of respiration is: Glycolysis + Kreb’s cycle = Glucose + 4ADP + 4H3PO4 + 8NAD+ + NADP+ +2FAD 54 . Oxidative decarboxylation of isocitric acid (a) dehydration and (b) decarboxylation) 5. 6. Dehydrogenation of malic acid in OAA. Condensation of Acetyl-CoA with oxalo-acetic acid 3.A 2. Oxidative decarboxylation of α-Keto glutaric acid. 7.{(a) dehydration and (b) hydration)} 4. Aerobic oxidation of P. Dehydrogenation of succinic acid into fumaric acid 8. Isomerisation of citric acid into isocitric acid . Conversion of succinyl CoA into succinic acid.\ Diagram of mitochondria All the chemical reaction of Kreb’s cycle can be summarized in following steps: 1. Hydration of fumaric acid into malic acid 9.

NADP. The pairs of hydrogen atoms released a series of coenzymes and cytochromes which form electron transport system. pyruvic acid. FAD etc.in this process ATP molecules are released (1NADH2 = 3ATP. enter into Kreb’s cycle for oxidation. Because two molecules of P. succinec acid and malie acid are oxidized. 2H 2H+ + 2e- These protons and electrons are accepted by various hydrogen acceptors like NAD.6CO2 + 4 ATP + 8NADH + 10H+ +2NADPH + 2FADH2 Thus as a result of oxidation of pyruvic acid.3. 2FADH2 = 4 ATP and two molecules of ATP are synthesized from 2GTP. before reacting with O2 to form H2O. 1FADH2 = 2ATP). isocitric acid. In the process of Kreb’s cycle 8 molecules of NADH2 =24ATP . a total of 6CO2 molecule will be evolved. one molecule of CO 2 in oxidative decarboxylation and two molecules of CO2 in Kreb’s cycle are liberated.A. 3. After accepting hydrogen atoms these acceptors get reduced to produce NADH2.4 ELECTRON TRANSPORT SYSTEM AND OXIDATIVE PHOSPHORYLATION Electron Transport System (ETC) During repiration simple carbohydrates and intermediate compounds like phosphoglyceraldehyde. 2PA * 3CO2 = 6CO2 All the NADH2 and FADH2 are oxidized to NAD and FAD through a chain of reaction c/a etc. The total number of CO2 evolved becomes 3 which indicates that 3 carbon pyruvic acid has been completely oxidized in glycolysis. ½ O + 2H+ + 2eH2O 55 . which are formed by one molecule of glucose in glycolysis. α− ketoglutaric acid. Each oxidative step involves release of a pair of hydrogen atoms which dissociates into two protons and two electrons. NADPH2 and FADH2.

Nicotinamide adenine dinucleotide (NAD). 3. Co-enzyme Q or ubiquinone. NADH2 or FADH2 produced by glycolysis and the TCA cycle. Specific enzymes of this chain receive electrons from reduced prosthetic groups. 7. 5. 4.S.2NADH + O2 + 2H+ 2NAD++ 2H2O As you know that H ions and electrons removed from the respiratory substrate during oxidation do not directly react with oxygen. The electrons are then transported successively from enzyme to enzyme. Cytochome-b 6. This process takes place in mitochondrial cristae which contain all the components of E. Mechanism of action of electron transport system: During respiration electron pairs liberated from respiratory compounds are accepted by coenzymes like NAD or NADP and FMN etc. Flavoproteins (FAD and FMN). 2.T. The transfer of electrons in all compounds except succinic acid takes place first in NAD+ or NADP+ and later on in FAD. The proteins of the inner mitochondrial membrane act as electron transporting enzymes. Fe-S protein complex. Instead they reduce acceptor molecules NAD and FAD to NADH2 and FADH2. 56 . The transfer of electgrons from succinic acid takes place diretly to the FAD and not through NAD+ or NADP+. All the above enzymes are found in F1 particles of mitochondria. Components of electron transport system: the electron transport system is made up of following enzymes and proteins: 1. Cytochrome-c 8. Cytochrome-a 9. They are arranged in an ordered manner in the membrane and function in a specific sequence. This assembly of electron transport enzymes is known as mitochondrial respiratory chain or the electron transport chain. Cytochrome-c1. These molecules then transfertheir electron to a system of electron acceptors and transfer molecules. down a descending ‘stairway’ of energy yielding reactions. Cytochrome-a3.

57 . Cyt-c. 2e. The electrons and proton are transferred to NAD causing its reduction and one proton is released in the medium. Cyt-a. 2H 2H+ + 2e(protons) (electrons) NAD + 2H+ + 2eNADH + H+ (reduced) (ion pool) 2. Cyt-a3 and then to oxygen atoms. are as follows: 1. The electrons pass to cytochromes Cyt-b.T.S. The hydrogen pair from succinic acid is first transferred to FAD to form FADH2. The energy from such electron transport is utilized in transporting protons from the matrix across the inner membrane to its outer side. The free energy released at this step is stored during oxidative phophorylation and one molecule of ATP is generated fronm ADP and inorganic phosphate. Oxygen atom accepts those electrons and reacts with hydrogen ions of the matrix to form water. Hydrogen pairs released from different substrates of Krebs cycle except succinic acid reacts with NAD+. This creates a higher proton concentration outside the inner membrane than in the matrix. The FADH2 transfers electrons to coenzyme Q throught Fe-S and CoQ.and one H+ are transferred from NADH to FAD causing oxidation of NADH to NAD and reduction of FMN into FMNH 2. At each step of electron acceptor has a higher electron affinity than the electron donor from which it receives the electron. The difference in proton concentration across the inner membrane is called proton gradient. One H+ is picked up from hydrogen ion pool to complete this reaction. Now. Different Steps of E.Due to this reason only two molecules of ATP are formed in the formation of fumaric acid from succinic acid whereas in case of other compounds 3 ATP molecules are produced because these cases the electrons are first picked up by NAD. Cyt-c1. O2 + 4e2(O--) + 4e+ 2(O--) 2H2O Oxygen is thus the terminal electron acceptor of the mitochondrial respiratory chain.

Each cytochromes possesses an iron elements in the centre which functions for accepting (Fe3+ Fe2+) or donating (Fe2+ Fe3+) When a cytochrome accepts electrons. When the electron transfer from cytochrome-b to cytochrome-c1. it is oxidized. ADP + iP O 2 ATP E. II. Just as a flow of water from a higher to lower level can be utilized to turn a water-wheel or a hydroelectric turbine. Since FADH2 donates its electrons further down the chain. III. When NADH2 is oxidized to NAD by reacting with FAD. protons return to the matrix down the proton gradient. Now it is clear that oxidation of one molecule of reduced NADH2 or NADPH2 results in the formation of 3 molecules of ATP while oxidation of FADH 2 leads to the formation of 2 molecules of ATP. it is reduced and if it donates electrons.T. Transport of two electrons from NADH2 by the electron transport chain simultaneously transfers three pairs of protons to the outer compartment. The return of proton occurs through the inner membrane particles. Therefore.The reduction of various cytochromes requires only electrons and no protons. During oxidative phosphorylation ATP molecules are produced during following steps: I. 58 . the energy released by the flow of protons down the gradient is utilized in synthesizing ATP. This process of synthesis of ATP molecules from ADP and inorganic phosphate by electron transport system of aerobic respiration called as oxidative phosphorylation. When the electron transfer from cytochrome-a to cytochrome-a3. Chain The process of oxidative phosphorylation takes place in mitochondrial crests through electron transport chain. In the F0-F1 complex the F1 head piece functions as ATP synthetase. Its oxidation can only produce two ATP molecules. oxidative phosphorylation produces three ATP molecules per molecules of NADH2 oxidized. Oxidative Phosphorylation In all living beings ATP generated during oxidative breakdown of complex food products. The latter synthesizes ATP from ADP and inorganic phosphate using the energy from the proton gradient. One high energy ATP bond is produced per pair of protons returning to the matrix through the inner membrane particles. Due to high proton concentration outside the inner membrane.

Total 36 ATP molecules are produced during whole process.3.P. Warberg et al (1935) & Dicken’s suggested an alternative oxidative pathway for glucose oxidation.3. Total 12 NADPH2 molecules are formed in each cycle which are oxidized by cytochrome cycle into 12 molecules of NADP. It is named as Pentose Phosphate Pathway or Hexose Monophosphate shunt. Out of 6 only molecule of glucose monophosphatic is oxidized into CO2 in each cycle of P. 59 .P &5 molecules of fructose monophosphatic & glucose monophosphatic are formed.5PENTOSE PHOSPHATE PATHWAY : AN ALTERNATIVE PATHWAY FOR GLUCOSE BREAKDOWN.

when glycolysis & Kreb’s cycle do not occur due to some reason. 1.P. Write your answer in the space given. 3. End product of anaerobic respiration is-------------------------4. Respiratory enzymes are found in ------------------------------2. 5-carbon compounds produced are used in the synthesis of nucleic acids. CHECK YOUR PROGESS (3) Note: 1. This pathway can supply required quantity of the energy to the cell. 2. Write elaborated form of following (a) NADP -----------------------------60 . It is substitute for glycolysis and Kreb’s cycle. Pace maker of glycolysis is---------------------------------------3. No. 2.Significance of P. Compare your answer with those given at the end of the unit 1.P. of ATP formed during respiration is -----------------------5.

A can not be generated from pyruvate or PEP.6 GLYOXLATE CYCLE Kreb’s cycle Acetyl CoA Glyoxylate cycle • • • It is special modification of TCA cycle It comes into play during oxidation of acitic acid or substrate (F. Value of R.A. at this time O.A) that are converted to acetyl CoA with out the intermediate formation of pyruvate Also.A.A .3. Like CH3 C=O + CO2 + ATP COOH Pyruvate Or CH2 C-O~P + ADP + CO2 COOH CH2 + ATP 61 COOH CH2 C=O + ADP + P COOH O.Q for fatty germinating seeds is ---------------------3. a mechanism available in case of anaerobes only.(b) TCA ---------------------------------(c) PPP---------------------------------(d) EMP -------------------------------6.

• • CO2 evolving steps of kreb cycle are bypassed Glucose or fattyacids can serve as source of acetate. precursor of OAA.A.lyase H – C – COOH OH – C – n COOH CH2 CH2 COOH Succinate Glyoxylate + COOH COOH H–C=O Isocitrate Regulation of enzyme: Pyruvate carboxylase • Enzyme has biotin as cofactor which acts as mobile carrier of CO2 in the of Mn++ ion. 62 . supply of O.A required for oxidation of acelate is replenished by the oxidation of succinate and malate which are produced through sequence of two reaction. The oxidation of pyruvate by pyruvate OH complex is completely irreversible. the twonreactions (succinate & malate formation) consitute a bypass where by two carbon atoms lost from TCA cycle are preserved as glycoxylate which than combines with acetyl CoA to form malate. • • Glyoxylate cycle acts as anapleurotic cycle allowing normal TCA cycle to function. In combination .COOH C=O COOH PEP • O.A In aerobic microorganisms there is no mechanism for synthesis of pyruvate from acetate. COOH CH2 I.A.

3.• Enzyme catalyses the reaction in which O. However. If ATP is less than OAA enters Kreb’s cycle.A + Biotin (enz) Acetyl CoA More over. alternative respiratory pathway branches from the cytochrome pathway at the ubiquinone pool in mitochondria. Biotin is not carboxylate unless acetyl CoA is not bound to enzyme • • A high level of acetyl CoA signals need for more O.A. In specific thermogenic floral tissues of some plants. because the alternative pathway can be active even when the cytochrome pathway is not saturated. a high rate of electron flow through the alternative pathway generates heat and volatilizes compounds that attract insect pollinators. it ATP is more than OAA is consumed in gluconeogenesis.7. Enzyme pyruvate carboxylase has allsoteric site for acetyl CoA is activated only in +nce of Acetyl CoA.A.A • • Biotin +ATP + HCO3Biotin~ CO2 + pyruvate (enz) Mn ++ Biotin ~CO2 + ADP + Pi O.Carboxylase Pyruvate CO2 & ATP O. Thus. ALTERNATIVE OXIDASE SYSTEM In all plants and many fungi. Electron flow through this alternative pathway is not coupled to ATP synthesis at two of the three proton translocation sites.A. the alternative pathway may allow continued turnover of carbon skeletons through glycolysis and the tricarboxylic acid cycle when the ATP concentration is high and inhibits ATP synthase activity so that the pathway can function as an energy overflow mechanism. In non-thermogenic plant tissues. it is likely that the two pathways are regulated in concert to balance ubiquinone 63 . a cyanide-resistant.3.A.A.A is formed from carboxylation of pyruvate P.

Based on assumptions about the enzyme's structure. and correspond to Cys-78 and Cys128 in the deduced sequence of the Arabidopsis thaliana alternative oxidase used in this study. These conserved Cys residues are the likely candidates for formation of the intersubunit disulfide bond and the site of -keto acid activation. but have less effect on the oxidized form of the enzyme. In the present study. was involved in the sulfhydryl/disulfide system. it was predicted that the more NH2-terminal Cys-78.pool oxidation/reduction and carbon skeleton turnover in response to cytosolic ATP levels. and Cys-128. notably pyruvate. Based on homology alignment of amino acid sequences deduced from cDNA sequences. use of sulfhydryl reagents showed that the site of -keto acid action also involves a sulfhydryl residue. was the site of -keto acid action. which may be exposed to the mitochondrial matrix. Recently. The alternative oxidase is a homodimeric protein in the inner mitochondrial membrane and the oxidation/reduction state of an intersubunit disulfide bond has been demonstrated to regulate its activity. The phenomenon of respiration resistant to cyanide is connected with 64 . the alternative oxidases of higher plants contain only two conserved Cys residues. A second mechanism of activation of the alternative oxidase involves -keto acids. The same study indicated that this sulfhydryl is different from the one involved in disulfide bond formation. which is believed to be located within the mitochondrial matrix before the first putative membrane-spanning helix. heterologous expression in Escherichia coli cells. Both occur in the NH2-terminal domain of the protein. probably through the formation of a thiohemiacetal by reaction between the cysteine sulfhydryl and the -keto acid. and subsequent cross-linking and activity assays to determine that the cysteine residues involved in regulatory disulfide bond formation and -keto acid stimulation are identical. which may lie closer to the postulated catalytic site near the membrane. we have used site-directed mutagenesis. Cyanide-resistant respiration was discovered at the beginning of the 20th century as a curiosity in thermogenic plants during anthesis and was later found to be a typical feature of plant respiration. which significantly activate the enzyme when the regulatory sulfhydryl/disulfide is in the reduced state.

Q.write your answer in the space given Answer the following Questions. 38 65 . mitochondrial matrix b. phosphofructokinase c. Check your progress -The key 3 a.the presence in the respiratory chain of an additional terminal oxidase — alternative oxidase (AOX).1 Write short notes on.2 Write a short note on ATP – the biological energy currency of the cell.. (a) Glycolysis (b) PPP (c) TCA (d) Glyoxylate cycle Q. ethyl alcohol d. CHECK YOUR PROGRESS (4) note.

pentose phosphate pathway. 1943. waxes. It is stored in special plastids known as elacoplastids. LIPIDS are esters of fatty acids and alocohol which form emulsions with water but are soluble in organic solvents. E & K . Tricarboxylic acid cycle. Adenosine di phosphate. and role. f. Parnas pathway. these are all lipids or rich in lipids. It includes fats. Vit A.e. energy liberated on hydrolysis. of substance like cooking oil. The main component of most of the lipids is fatty acid. this term lipid was for the first time used by Blour. You are familiar with a no. OBJECTIVE : 66 . Lipids are soluble in organic solvents like alcohol and ether and insoluble in water. R – COOH + HO – C2H5 R – C2H5 + H2O + CO2 Fats and their derivatives are collectively called lipids.less than one. In plants. Check your progress-the key-4 Your answer must include 1An outline of cycles and names of important enzymes 2structure of ATP.4. glycolipids and sterols. Nicotinamide adenosine diphosphate.LIPID METABOLISM_______________________________________________ INTRODUCTION : In this unit you will learn about structurally distinct organic compounds called lipids. 3. butter. lipids are found in their seeds and fruits. Embeden Mayernoff. natural rubber and cholesterol. Greek – lipose – Fat. phospholipids. menthol and Eucalyptus oil are also fats. waxes. Plants pigments like carotene of carrots and lypocene in tomatoes.

STRUCTURE AND FUNCTIONS OF LIPIDS STRUCTURE Chemically. 2. An understanding of the biosynthesis process of important lipids in plants.4. Understand the basic component of lipids.After studying this unit you will be able to: 1. 4. lipids are the esters or glycerides of fatty acids and glycerol. 6. Structure and functions of lipids. During the formation of Lipids. Various types of lipids that occur in nature. the carboxylic group (-COOH) of each fatty acid react with alcoholic group (-OH) of glycerols to form a esterlinkage. 3. And how lipids are broken down to liberate vast amount of energy.A 3 molecules + HO – CH2 + HO – CH2 + HO – CH2 glycerol CH3 (CH2)nCOOCH2 CH3 (CH2)nCOOCH CH3 (CH2)nCOOCH2 Triglycerides 67 .1. CH3(CH2)nCOOH CH3(CH2)nCOOH CH3(CH2)nCOOH F. R – COOH Fatty Acid + HO – C2H2 Alcohol R – C2H5 + H2O + CO2 When lipid is made of three molecules of FA and one molecule of glycerol it is called as esterlinkage. Role played by lipids in living beings specially in plants 5. 3.

All the FA have a long chain of hydrocarbon. 68 . R2.When all the fatty acid acyl group or R1. Thus the nature and structure of lipids depends upon the nature and structure of their FA. & R3 group are identical then this product is called as simple glyceride. When it contains different kinds of FA acyl group it is called mixed triglycerides.

g..Sulphatides Phosphoglycerides e..g. Simple lipids Compound lipids Fatty acids Derived lipids Fats and oils e.g.General classification of lipids. Sphingomyelins Terpenes e. Inocitol Carotenoids Phosphosphingosides e.g. Carotenes. Bee wax Aclohols Phospholipids Glycolipids e.g. Hormones 69 . cephalins Proteolipids e.g. lecithins. Lycopene e.g. Kerasin. Triglycerides Waxes e.g.g.g. Nervon Sulpholipids e.. Cholesterol. Lipoproteins Steroids Phosphoinositides e.

position of bonds.F. 2. differ from one another in chain length. Only over 100 F. F.linoleic acid. Possess a long hydrocarbon chain and a terminal carboxy group. _ eg:-OOC (saturated) CH3 C16 Palmitic acid. of. F.FATTY ACIDS Essential FA: Mammals can synthesize saturated and monosaturated F. occurs in large amounts as building block components of complex lipids. ốcc C18 Oleic acid (unsaturated) OOC CH3 Saturated F. from other precursors but are unable to make linoleic acid & γ. are required for synthesis of prostaglandins – which are hormone like component and are required in trace amounts in many physiological conditions.A. no. have been isolated from cells to occur in free or unesterified forms. All Fatty acids 1.A (monoenoic) 70 .A.A. Unsaturated Polyunsaturated F.A.A (polyenoic) Monounsaturated F. There bore : obtain them from plant source.A.A. carbon chain may be saturated or unsaturated 3. E.

In higher plants and animals living at lower temperature Unsaturation α fluidity α 1/ temperature At lower temperature fluidity increases. called as octadecanoic acid because parent is octadecane • • • • A C18 F. • The systematic name for a fatty acid is derived from the name of its parent hydrocarbon by the substitution of oic for the final e.A.A.10 double bond and methyl terminal end.A. Marine animals have odd carbon F.A.A. Bacteria contain fewer types and simpler F.A. carbon atoms are numbered starting at carboxy terminus. 6. with one double bond is called as octadecenoic With two bond – octadecadienoic acid With three bond . Viz C12 to C18 saturated F.A. C18 sautrated F. F. than higher organisms (obviously). Also in most poly unsaturated F.: in most monounsaturated acids double bond occurs between carbon no. 3. Nomenclature of F. F.A is . to keep organism working no. in poly unsaturated F. b) Unsaturated F. first double occurs between carbon 9&10 and others between 9. of saturated fats increases. NOTE: 1. F.A.octadecatrienoic acid. Terrestrial animals have only trace amounts of odd carbon no. 5. the double bonds are separated by one methylene group. The double group in all naturally occurring saturated F. 9 & 10.A. Unsaturated F.A.A.A. predominate over saturated ones. 4. 71 e.a) Saturated F. are CIS geometrical configuration only a few are trans.A in abundance.A C16 & C18 monosaturated F. 2. with more than one double bonds have not been found in bacteria. have higher melting point and are liquid at normal temperature.A.g.: Do not have double bonds.

Saturated and unsaturated F. coconut and castor the fat is stored by the plants to provide nourishment for the embryo during germination. bromine gives information about number of double bonds therefore they undergo addition reaction.hydrocarbons tails are flexible and can exists in very large number of coformations because each single bond has freedom of rotation.A. 12:0 14:0 16:0 16:0 18: CH3(CH2)10COOH CH3(CH2)12COOH CH3(CH2)14COOH N-dodecanoic Laurica N-tetradecanoic Myristic N-hexadecanoic Palmitic Unsaturated F. a) In saturated F.ω H3C • • • (CH2)n 3 β CH2 α CH2 2 C 1 O n Carbon atom 2 & 3 are often referred as α and β respectively Methyl carbon at distal end is known as ωcarbon The position of double bond is represented by symbol followed by a subscript no.A. 1.A show one or more rigid ‘Kinks’contributed by non rotating double bond(s).A. Oil extracts from these seeds is used for cooking 72 . CH3(CH2)6CH=CH(CH2)7COOH Palmitoleic CH3(CH2)7CH=CH(CH2)7COOH Oleic acid Physical and Chemical properties of F.A on quantitative titration with halogens iodine.A. c) The CIS conformation of double bond produce a bend in aliphatic chain whereas trans resembles saturated extended conformation. have quite different conformations.A : Saturated F. mustard. d) Unsaturated F. FUNCTIONS: 1) Food reserve: In oil seeds such as groundnut.A. Some naturally occurring F. b) Unsaturated F.

saturated and unsaturated fatty acid occurs in nature different pathways are followed for synthesis of different F. 1) Biosynthesis of saturated F. two types of fatty acids Viz .A.D& E.4. here we will learn anabolism/biosynthesis of fatty acid. 1.2) Structural Constituents: lipids are involved in building of cellular components like cell membrane. 5) Fat transport: phospholipids place an important role in absorption and transport of F. FATTY ACID BIOSYNTHESIS As you know metabolism involves both anabolism and catabolism. Lipids are esters of -----------------------------------------------2. 73 . Oxidation of fatty acids takes place in ------------------------3.2. CHECK YOUR PROGRESS (5) Note: 1. Since. (malonyl CoA pathway) 2) Enzymatic biosynthesis of unsaturated F.A. Give two examples each of --------------------------------------(a) simple lipids ---------------------------------------------------------------(b) Saturated fatty acids & unsaturated F. ----------------------------------------------------------------------------------------------------------------------------------- LIPID METABOLISM 3.A -------------------------------(c) Functions of lipids.A. write your answers in the space given 2. 4) Harmone synthesis: cholesterol is most important sterol in animals. nuclear membranes and membrane of cell organelles 3) As solvent: lipid acts as solvent for the fat fat soluble vitamins like A. Check your answer with the one at the end of the unit.A.

Mn++ .CoA Carboxylase CH3 – CO – SCOA+CO2+ATP Mn++ (Acetyl – CoA)(2C) 74 Acetyl-CoA + ADP+Pi + OAA . In fatty acids(participating in fat synthesis) the number of carbon atoms varies form 16 to 18. Biotin – Acetyl .Biosynthesis of Fatty Acids (i) Biosynthesis of saturated Fatty Acids: Naturally occurring fatty acids may be saturated and main pathway of biosynthesis of fatty acid in plants.ADP and inorganic phosphate. According to Green (1960). Before starting of fatty acid biosynthesis an important preparatory reation. Two enzymes complexes and five cofactors-ATP. NADPH and CO2 are essential for the synthesis of fatty acids. Ultimate source of carbon atoms of fatty acid is acetyl-CoA which is produced from carbohydrates and aminoacids. The long chain compounds of fatty acids are synthesized from two carbon compounds Acetyl-coenzyme A (Acetyl-CoA) which is highly reactive compound and is produced as an intermediate in respiration of sugar and fats. Acetyl-CoA is regenerated with the help of citrate cleaving enzymes as follows: Citrate Cleaving Citric Acid + CoA +ATP Enzymes Acetyl-CoA acts as a primer. Molecules of malonyl-CoA are successively attached to the primer molecules of Acetyl-CoA accompanied by decarboxylation. the formation of malonyl-CoA takes place in cytosol. biotin. acetyl-CoA react with CO2 and ATP to produce malonyl-CoA. The synthesis of fats takes place in stepwise reaction taking place again and again. 2-carbons atoms of acetyl-CoA are added in the chain.Complete biosynthesis of fatty acid takes place in cytosol. Overall reaction is catalysed by the complex of 7 proteins-the fatty acid synthetase complex. In presence of biotin-acetyl-CoA carboxylase enzyme. In each step. animals and bacteria is common and takes through malonyl CoA pathway. Mn++ acts as cofactor in this reaction.

CO2 is released during this reaction and water (H2O) and NADP are formed. H | | O || H O | | 75 || Acetyl-Coa(2C) O || H__ C__ C__ S. acetomalonyl-CoA is con verted into 4 carbon compound butyryl-CoA. H | | O==C__ OH Malonyl-CoA(3C) HO H O | || | | | || H__C__C__C__C__S.CoA+CH3___CO___ S.CoA+ADP+iP O == C__ OH Malonyl-CoA(3C) Malony I-CoA react with acetyl-CoA to produce acetomalonyI-CoA (5C compound).H | | O || H__ C__ C__ S.CoA+ 4NADPH .CoA H__ C __ C __ C __ C __ S.CoA H O=C_ OH Acetomalonyl-CoA(5C) In presence of specific enzyme and coenzyme NADPH.

4. 3. SYNTHESIS OF MEMBRANE LIPIDS Biological membrane A biological membrane or biomembrane is an enclosing or separating amphipathic layer that acts as a barrier within or around a cell. (ii) Biosynthesis of unsaturated fatty acid: Biosynthesis of unsaturated fatty acid has not been completely studied in higher plants. almost invariably.H O==C__ OH Acetomalony-CoA(5C) H | | H H || | H | H H O | || H__ C __ C __ C __ C __ S. When the chain length reaches 16 or 18-carbons.CoA+CO2 +4NADP=H2O Butyryl-cOa(4C) The butyryl-CoA then reacts with another molecule of malonyl-CoA and cycle is repeated and 6-carbon compounds is produced. with occasional proteins intertwined. This will again react with still another molecule of malonyl-CoA to produce 8-carbon compounds. some of which function as channels. Most 76 . double bonds are introduced into previously formed saturated fatty acid by the enzyme desaturase. a lipid bilayer. For example.Such membranes typically define enclosed spaces or compartments in which cells may maintain a chemical or biochemical environment that differs from the outside. composed of a double layer of lipid-class molecules. Enzymes acyl-CoA desaturase and stearyl-CoA desaturase have been isolated from yeast and Euglena respectively. specifically phospholipids and cholesterol.3. It is. and the plasma membrane separates a cell from its surrounding medium. the fatty acid is released. During their synthesis. the membrane around peroxisomes shields the rest of the cell from peroxides.

The heads of phospholipids are phosphorylated and they consist of either: Glycerol (and hence the name phosphoglycerides given to this group of lipids). it could either go through one of the protein channels or be taken in by means of endocytosis. The three major classes of membrane lipids are phospholipids. and other chemical properties of the atoms and molecules attempting to cross it will determine whether they succeed to do so. Sphingosine (with only one member . 77 . Selective permeability is essential for effective separation of a cell or organelle from its surroundings. glycolipids. nonpolar (hydrophobic) hydrocarbon chains linked to a hydrophilic head group. Probably the most important feature of a biomembrane is that it is a selectivelypermeable structure. Phospholipids Phospholipids and glycolipids consist of two long. and are called membrane-bound organelles. charge.organelles are defined by such membranes.sphingomyelin). If a particle is too large or otherwise unable to cross the membrane by itself. Biological membranes also have certain mechanical or elastic properties. This means that the size. but is still needed by a cell. and cholesterol.

in the case of sphingomyelin and the glycolipids.and glycolipids usually contain an even number of carbon atoms. FAs may be saturated or unsaturated. with the configuration of the double bonds nearly always cis. Fatty acids The fatty acids in phospho. The length and the degree of unsaturation of FAs chains have a profound effect on membranes' fluidity. The 16. 78 .and 18-carbon FAs are the most common ones. one FA and the hydrocarbon tail of sphingosine .in the case of the phosphoglycerides.Glycolipids The heads of glycolipids contain a sphingosine with one or several sugar units attached to it. The hydrophobic chains belong either to: two fatty acids . typically between 14 and 24.

Flow chart for the biosynthesis of fatty acids. triglycerides and cholesterol are as follows: FATTY ACID BIOSYNTHESIS 79 .


81 . A phospholipids molecule is a bipolar. represent the water repellant hydrophobic end and the phosphate containing end is the hydrophilic end. It is the chief constituent to cell membrane. sulphur or protein. the non-polar tails. Its two long fatty acid molecules . metabolism and blood clotting.4. COMPOUND LIPIDS Compound lipids contains some additional groups or elements besides fatty acid and alcohol. It helps in transport. 1) Phospholipids: These lipids contains phosphoric acid beside alcohol and fatty acid. The additional group may contain phosphorus. Phospholipids include lecithins.CHOLESTEROL BIOSYNTHESIS 3. Phospholipids are present in all cells and control the permeability of membrane. STRUCTURAL LIPIDS & STORAGE LIPIDS.4. cephalinsand plasmomalogens. nitrogen.

Cephalins contains choline and sometimes serine in place of choline. linolinic and arachidonic acids are commonly found in lecithins. Lecithins and other phospholipids are yellowish-grey solids soluble in ether and alcohol but insoluble in acetone. • Cephalins: Cephalins are found in animals tissues in close association with lecithin. The difference between the lecithins and cephalins is in the nature of nitrogenous base. On exposure to air. linoleic and arachidonic acids • Plasmalogens : Plasmalogens are abundant in brains and muscles. fatty acid. Their structural formula is as follows. The fatty acid components of cephalins are usually stearic. oleic. These lipids are soluble in all lipids solvent. Palmitic. choline. stearic. phosphoric acid. Choline Phosphate Polar head group Glycerol Fatty acid 1 Fatty acid 2 Phospholipid 82 . In this group of phospholipids a complex aldehyde is attached to the α-carbons atom of glycerol. Ethanolamine or choline is attached with phosphoric acid. lecithinaseis present in venom of bee and cobra. Lecithins are broken down by the enzyme lecithinase to lysolecithin. they rapidly darken in color and absorb water forming dark greasy mass. They are also found in soyabean oil. oleic. They are found in the seeds of higher plants.• Lecithins: Lecithins are widely distributed in nature and on hydrolysis they yield glycerol.

They are found in cell membranes and other cellular components containing lipids.serocine. These are solid wax like substances. It contains sulphur and amino acids besides fatty acids and glycerol. Sterols are widely distributed in plants. Derived lipids include hydrolytic product of lipids and in addition other lipid like compounds such as sterols. carotenoids. It is insoluble in water. aldehydes.g. The best known animal sterol is cholesterol which is present high concentration in nervous tissue and in bile. animals ans micro-organisms.estrabiol and cortisol and vitamin D.O || Saturated O __H2C__O__C__R1 fatty acid || Unsaturated Fatty acid | O CH3 CH3 CH3 | H2C__O__P__OCH2__CH2_N | (H2OH)O Lecithin 2) Glycolipids: these lipids contain nitrogen and carbohydrates beside fatty acids.like progesterone. 3) Chromolipids : It includes pigmented lipids eg carotene 4) Aminolipids: It is also known as sulpholipids. frenocine. hydrocarbons. spleen and yolk of an egg e. alcohols. essential oils. Cholesterol is also the precursor of hormones.so it gets R2 __C__O__C__H 83 . They are highly soluble in fat solvent.testosterone . They are essential for plant growth and flowering. DERIVED LIPIDS Lipids obtained by the hydrolysis of simple and compound lipids are called as derived lipids. Cholesterol is a crystalline solid with rhombic crystals. The sterols are composed of fused hydrocarbon rings and a long hydrocarbon side chain. It is genrally found in white matter of nervous system. etc. ketones. Chemically they are alcohols and occur ether as such or as ester of fatty acid.

During this process triglycerides react with water (H2O) to produce fatty acids and glycerol. During seed germination. 3. Degradation or oxidation of fats involves following three processes: 1) Hydrolysis of fat into glycerol and fatty acids 2) Metabolism of glycerol 3) Oxidation of fatty acids. Hydrolysis of Fat into Glycerol and Fatty Acids Degradation of fatty acids starts with their hydrolysis. Some are straight chain carbon compounds without and side branches.deposited in the arteries and veins and hence .this may lead to high blood pressure and heart diseases. Ergosterol is present in food in small quantity. The whole process is completed in three steps. It is precursor of another form of vitamin D. It is found in argot and mould like yeast.lemon. blood cholesterol rises. Most essential oils are either terpenes related to isoprene or isopantene or benzene derivatives.etc. The various steps are as follows: 84 .eucalyptus. It is found as a result of reduction by bacteria in the intesting from the couble bond of cholesterol between C5 & C6. ergocalciferol (D2). OXIDATION OF LIPIDS OR DEGRADATION OF FATS. Essential oils are usually obtained from plants such as pine. pipermint.5. Essential oils are also grouped under lipids because of their solubility and natural occurrence. The enzyme lipase first attacks the α-carbon and then in the β-carbon atom.4. Coprosterol is found in faeces. the enzyme lipase catalyses this reaction. rose. Lipid is a storage material and can b used as energy rich fuel by cells during seed germination.

O.OH CH2OH Glycerol + R” – C .O.O. 85 .C – R” OH O αCH2.OH Step (1) and (2) are reversible and step (3) is irreversible.C – R”’ α-β-diglyceride (3) CH2OH O CH. Lipase H2O α-β-diglyceride αCH2OH O O βCH.C – R” + R”’ – C - αCH2OH β-monoglyceride Fatty Acid O CH. Lipase CH2OH Ca++.H Triglyceride (2) αCH2OH O βCH.C – R” CH2OH β-monoglycerid H2O Ca++.C – R2 O CH2OC – R3 Fatty Acid + R1 – C .O.(1) O CH2O – C – R1 O CHO – C – R2 O CH2O – C + H2O Lipase Ca++ – R1 CH2OH O CHO.

P 86 . the metabolism of glycerol takes place according to following diagram(Figure 13. which may be oxidized into Krebs cycle to CO2 and H2O Sugar Dihydroxyacetone Glycerol Pi α.glycerophoshate Glycolysis Pyruvic Acid TCA cycle CO2 + H2O Metabolism of glycerol Glycolysis Dihydroxyacetone . enters the carbohydrates metabolism and metabolized into CO2 and H2O through various steps. After complete metabolism. glycerol is converted into acetyl-CoA.9). According to Stumpf (1955) and Beevers (1956).2) Metabolism of Glycerol : Glycerol is produced during hydrolysis of triglycerides.

oxidation . On the basis of α. Derived lipids 3)Oxidation of Fatty Acids: The oxidation of fats depend upon α.and βcarbon atom. This type of oxidation is found only in the fats in which number of carbon atoms is limited from 13 to 18 carbons.oxidation of fatty acids is discovered by Newcomb and Stump in 1952. Compound lipids 2. (ii) β.CHECK YOUR PROGRESS (6) Write short notes on.or β-carbon atom.OXIDATION OF FATTY ACIDS α.oxidation. peroxidative decarboxylation of fatty acids takes place. In this oxidation only one carbon atom is removed in every step. Decarboxylation: First of all in the presence of fatty acid peroxidase enzyme. It is called α-oxidation because in this process oxidation of α-carbon atom takes place. * α-oxidation of fatty acids takes place only in cotyledons and young leaves. α. Mechanism of α-oxidation The α-oxidation is completed in following two steps: a. In this 87 . 1. the oxidation of fatty acids is of the following kinds: (i) α.

Peroxidase R-CH2.and C18)are utilized as substrate in α-oxidation.process fatty acids react with hydrogen peroxide(H2O2) to yield an aldehyde (one carbon atom shorter than fatty acid) CO2 and H2O. The acids with more than C12(usually C14.CH2-COOH + H2O2 α-carbon fatty acids R-CH2-CHO + CO2 + H2O Aldehyde Hydrogen Peroxide b. Dehydrogenase R-CH2-CHO + H2O Aldehyde NAD+ NADH + H+ This resulted acid is again utilized by the enzyme fatty acid peroxidase as substrate for another turn round the two stage of α-oxidation spiral. Dehydrogenation : the enzyme aldehyde dehydrogenase now catalyses the oxidation of aldehyde(formed by first enzyme)to yield the corresponding acid.C16. The enzyme of α-oxidation are specific for long chain saturated fatty acids. 88 . R-CH2-COOH Acid *The fatty acids formed after dehydrogenation may also be oxidized through β-oxidation.

CHO (2) ˚ etc.CH2CHO (1) = fatty Acid peroxidase .CH2CH2COOH H2O2 ˚ * H2O2 (1) H2O NADP + H+ NAD+ R. This is called β-oxidation because βcarbon (i. β-oxidation is the chief process of fatty acids degradation in plants.C-3) of the fatty acid is oxidized during this process. (2) = aldehydrogenase α-oxidation in peanut cotyledons β-OXIDATION OF FATTY ACID According to Knoop.COOH H2O (1) R.e. Requirements of β-oxidation β-oxidation requires the following substance: 89 . CO2 CO2 ˚ R.CH2COOH ˚ R. β-oxidation takes place in mitochondrial matrix (and also in glyoxyzones) and involves sequential removal of 2-C in the form of acetyl-CoA molecules from the carboxyl and of fatty acids. degradation of fatty acids takes place by successive removal of C2 units after oxidation of the β-carbon atoms.˚ * + R.

iii. ii. 90 .a) A fatty acid b) An energy source. Activation and entry of fatty acid into mitochondria i. Activation of fatty acids into mitochondria.ATP c) Coenzyme-A d) A carrier molecule – carnitine e) five enzymes: 1) Acetyl-CoA synthetase 2) Acetyl-CoA dehydrogenase 3) Enoyl-CoA hydrase 4) β-hyrdoxy acyl CoA dehydrogenase 5) Thiolase Mechanism of β-oxidation : Fatty acids are formed in cytosol. Transfer to carnitine. how fatty acids enter mitochondria for their degradation: These are the following steps by which fatty acid enters mitochondria: a. Transfer to intramitochondrial membrane. Now the question is.

In this reaction esterification of fatty acid takes place.SCoA || O 91 + AMP+PPi . CoASH is consumed and CoA derivative of fatty acid is produced. three steps: (a) OXIDATION OR DEGRADATION OF FATTY ACID 1) Activation and entry of fatty acids into mitochondria: It is completed in following Activation of fatty acids: the first step involves the activation of fatty acid in the presence of ATP and enzyme thiokinase.CH2.b.CH2COOH+CoASH+ATP ============== RCH2CH2C.Mn++ R. Thiokinase.

Fatty acyl-CoA Pyrophosphate (b) transfer to carnitine: the acyl group of fatty acid CoA is transferred to carnitine. In this step acyl group of fatty acyl carnitine is transferred to intramitochondrial CoaA. CH3 R__ CH2. which requires Acyl-CoA as substrate.CH2 C || O Fatty acyl-CoA SCoA+H O__ CH__CH2__N+ | CH2COO_ Carnitine Acyl-CoAcarnitine transferase CH3 R__ CH2.CH2 C__O__ CH__CH2__N+ || O | CH2COO_ CH3 + COASH CH3 CH3 CH3 Fatty acyl-CoA Carnitine (C) Transfer to intramitochondrial membrane: Dergadation of fatty acid takes place in mitochondrial matrix. CH3 R__ CH2.CH2 C__O__ CH__CH2__N+ || O | CH2COO_ CH3 + COASH CH3 Fatty acyl-CoA Carnitine 92 . Carnitine s a carrier protein which is found in inner mitochondrial membrane which transport fatty acyl CoA through inner mitochondrial membrane to the actual site of degradation.

β-hydroxyacyl-CoA in the presence of Enoyl hydrase. first of all two hydrogen atoms are reomoved from α.and β-carbon atoms of fatty acyl-CoA and trans – α.Acyl-CoAcarnitine transferase CH3 R__ CH2. Enoyl Hydrase R-CH=CH-CO-S-CoA +HOH R-CH(OH)-CH2-CO-S-CoA 93 . β α Acyl CoA dehydrogenase β α R – CH2-CH2-CO-S-CoA + FAD R-CH=CH-CO-S-CoA + FADH2 Trans α. β-unsaturated fatty acyl-CoA is formed.CH2 C__S__ CoA+OH__CH__ CH2__ N+ || O | CH2__COO_ Carnitine CH3 CH3 Fatty acyl S.β-unsaturated fatty acid b) Hydration: In the second step the addition of water molecule takes place to form . This reaction is catalysed by FAD containing enzyme acyl-CoA dehydrogenase.CoA 2) Oxidation or degradation of fatty acid : The oxidation of fatty acid involves in the following four steps: (a) First dehydrogenation : During oxidation.

palmitic acid enters seven times in the β-oxidation pathway for their complete oxidation. The thioclastic cleavage of β-keto fatty acyl-CoA takes place in the presence of enzyme β-keto acyl-thiolase and results in the formation of an active 2-C unit actyl-CoA and β-keto fatty acyl-CoA molecule which is shorter by 2-C atoms then when it entered the β-oxidation spiral.β -Hydroxy fatty acyl. The sequence continues until whole molecule is degraded.fatty acyl-CoA is unstable and it releases 2-C fragment as actyl –CoA by the process of thiolysis. β.hydorxy acyl-CoA dehydrogenase. *Acetyl CoA is used in TCA (=Krebs cycle) and Keto acyl-CoA reenters into β-oxidation for its complete oxidation and in every step two carbon atoms are released. C16H32O2 + 23O2 16CO2 + 16H2O 94 .CoA c) Second dehydrogenation: Now β-hydroxy acyl-CoA is dehydrogenated in the presence of NAD specific β. it is passed through β-oxidation pathway seven times and get completely oxidized to form CO2 and H2O. e.g.hydorxy acyl-CoA R-CH(OH)-CH2-CO-S-CoA + NAD+ Dehydrogenase R-CO-CH2-CO-S-CoA + NADH +H+ β-keto fatty acyl-CoA d) Thiolysis : β. which now bears a carboxyl function and β-keto fatty acylCoA is formed. Energetics of β-oxidation For complete oxidation of palmitic acid. Two hydrogen atoms are removed from β-C atom (βoxidation).

The efficiency can be calculated 130*8000*100 95 16 CO2 + 8H2O + 8CoA .g.Palmitic acid *Each turn of β-oxidation pathway produces 5ATP molecules. * Each acetyl CoA molecule after complete oxidation through TCA cycle produces 12 ATP molecules.A. one ATP molecule is utilized in activating the fatty acid molecule. Here 32 atoms of O2 are used. however the first turn shows a net gain of only 4 ATP.C. Thus .C. T. Their reoxidation takes place through ETS Step 1 : Palmitic acid 8 actyl –CoA +14 electron pairs 7 pairs of electrons via FAD+ 7*2 = 14 ATP 7 pairs of electrons via NAD+ 7*3 = 21 ATP 35 ATP 1 ATP used in activation of fatty acid – 1 ATP Net 34 ATP Step 2 : Now Acetyl-CoA enters the T. For e. 2. hence. cycle 8 Acetyl-CoA + 16 O2 Therfore 32*3 = 96 ATP formed.A cycle and oxidized. As we know that 3 ATP molecules are formed from each O2 atom during oxidative phosphorylation. Thus total ATP formed will be 34 + 96 = 130 ATP molecules gained. FADH2 and 1 NADH2 are formed in each cycle.: one molecule of palmitic acid (C16) after complete oxidation to CO2 and H2O produces 130 molecules of ATP as follows: 1. the total number of ATP molecules produced by a fatty acid depends upon the number of carbon atoms present in that fatty acid molecule. During activation 1 ATP is udes and 8 energy rich acetylCoA are formed.

1 what are plant lipids? How are they biosynthesized? Q. write your answers in the space given below. 96 .2 where in the cells does synthesis and breakdown of fats occurs? Q.= 2340.mitochondria and glyoxysomes 3.3 why does a gram of fat yields more energy than a gram of carbohydrate? Q.4 name the main steps and enzymes of B-oxidation pathway? Check your progress(5) – The Keys 1.Check your answer with the one at the end of the unit. Q.000 = 49% The remaining energy is lost in the form of heat. 2. Fatty acids 2. (a) Triglycerides and Bee wax (b) Butyric acid and caproic acid. Oleic acid and linoleic acid (c) Food reserve and Hormone synthesis. Answer the following questions in short in the space given below. CHECK YOUR PROGRESS (7) Note: 1.

mitochondria. Where do they occur in cell. Names of enzymes and types of each reaction.Check your progress – (6) Your answer must cover following points. It is the unique ness of this process to convert solar energy into chemical form. • • • • • • Structure of compound and derived lipids. SUM UP:PHOTOSYNTHESIS → Photosynthesis is the most significant process on earth for survival of life forms. Three main steps of lipid biosynthesis.A & lipids Synthesis occurs in cytosol Out line of oxidation pathways. Check your progress – (7) Your answer must cover following points. This conversion occurs in specialized structures in specialized structures in plants called as chloroplast T → The light reaction occurs in two ways a) Cyclic photophosphorylation b) Non-cyclic photophosphorylation → Light reaction is also called as Hill reaction. glyoxysomes. An out line classification of lipids. Outline of synthesis of F.4. 2. Oxidation occurs in cotyledons. 3. There functions Pathway of α -oxidation It occurs in cotyledons and young leaves. 1. 3. young leaves. 97 .6. B-oxidation pathway.

→ Dark Reaction/ Calvin cycle is signified for carbondioxide fixation. → Cytoplasm & Mitochondria are sites of respiration. a. → Photorespiration is a wasteful activity of the enzyme. b. → Phosphoglyceric acid is the first stable molecule formed during the process. 2. It leads to generation of reduction power. → All together 18 ATP and 12 NADPH2 are consumed in CO2 fixation. → CAM cycle is another pathway for CO2 fixation in Succeulent plants.→ Photolysis of water occurs in photosystem –II → O2 evolved comes from water. → The process of respiration is significant for plant because 1. → RUBISCO. 98 . Oxidation of pyruvic acid c. It leads to generation of precursors. → C4 cycle is an alternative pathway of CO2 fixation. → Process is completed in three steps. ETC & oxidative phosphorylation → Glycolysis is a fermentative pathway.Ribulose bis phosphate carboxylase is the main enzyme of cycle. → This enzyme fixed CO2 at higher concentration and is also responsible for photorespiration at higher concentration of oxygen. ATP is generated through it. → Phosphofructokinase enzyme is the pacemaker of pathway. 3. RESPIRATION → The process of respiration is an oxidation-reduction process. → ATP and NADPH2 are generated during light reaction. with OAA as first stable product. Glycolysis/EMP pathway. → 8 ATP are generated during oxidation of glucose to pyruvic acid. → Kreb’s cycle occur in mitochondria → All the enzymes of TCA cycle are located in inner mitochondrial matrix. → Plants perform aerobic reaction.

b. ASSIGNMENT 1.→ All together 38 ATP are generated through glycolysis through glycolysis and TCA cycle. CAM Pathway c. → Oxidation of Fatty Acids occurs though α & β . → Pentose phosphate pathway leads mainly to generation of reducing power (12NADPH2) and carbon required for nucleic acid synthesis. Draw and Explain Glycolysis is a fermentative pathway? 5. What are lipids? Give general classification? 7. glyoxysomes. Draw well-labelled Flow Charts of? a. Saturated F-A is synthesized through tmalonye COA pathway. mitochondria. → Glyoxlated cycle is a special modification of TCA cycle & is an anapleurotic cycle. → Fatty Acids are saturated or unsaturated type. Draw and Explain (a) Glycolysis (b) TCA (c) PPP 6. compound lipids and derived lipids. 3. → Oxidation occurs in cotyledons. Calvin Cycle.oxidation in stepwise enzyme mediated process. 3. Photorespiration d. What is the significance of light reaction in photosynthesis? 4. → ETC is located in inner membrane of mitochondria.4. Electron Transport Chain. 99 .7. Write a detail note on lipid metabolism. LIPID METOBOLISM → Lipids are esters of fatty acid and glycerol/ alcohols → They are broadly classified as simple lipid. Explain oxygenic photosynthesis? 2. → For synthesis of F-A different pathways are followed a. young leaves. Unsaturated F-A is synthesized through enzymatic pathway. b.

N. Sinha. 6. Kalyani Publication 7.4. Biochemistry.8. REFERENCE 1. Devlin & Witham CBS Publishers & Distributers. Pandey & B.H.A. Stryer.3. Berry. 100 .K. J. Osmund & G. Botany. S. A. C. Salisbury & Ross. 5. 2. Dutta.B. Plant physiology DBS publishers and Distributers. Lehninger. Lorimer Plant Physiology.C. Biochemistry. 4. Vikas Publication 3..

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