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# BIOMOLECULES Plant cell walls are made of cellulose. Paper made from plant pulp and cotton fibre is cellulosic. There are more complex polysaccharides in nature. They have as building blocks, amino-sugars and chemically modified sugars (e.g., glucosamine, N-acetyl galactosamine, etc.). Exoskeletons of arthropods, for example, have a complex polysaccharide called chitin. These complex polysaccharides are mostly homopolymers. # 9.6 NUCLEIC ACIDS The other type of macromolecule that one would find in the acid insoluble fraction of any living tissue is the nucleic acid. These are polynucleotides. Together with polysaccharides and polypeptides these comprise the true macromolecular fraction of any living tissue or cell. For nucleic acids, the building block is a nucleotide. A nucleotide has three chemically distinct components. One is a heterocyclic compound, the second is a monosaccharide and the third a phosphoric acid or phosphate. As you notice in Figure 9.1, the heterocyclic compounds in nucleic acids are the nitrogenous bases named adenine, guanine, uracil, cytosine, and thymine. Adenine and Guanine are substituted purines while the rest are substituted pyrimidines. The skeletal heterocyclic ring is called as purine and pyrimidine respectively. The sugar found in polynucleotides is either ribose (a monosaccharide pentose) or 2’ deoxyribose. A nucleic acid containing deoxyribose is called deoxyribonucleic acid (DNA) while that which contains ribose is called ribonucleic acid (RNA). # 9.7 STRUCTURE OF PROTEINS Proteins, as mentioned earlier, are heteropolymers containing strings of amino acids. Structure of molecules means different things in different contexts. In inorganic chemistry, the structure invariably refers to the molecular formulae (e.g., NaCl, MgCl, etc.). Organic chemists always write a two dimensional view of the molecules while representing the structure of the molecules (e.g., benzene, naphthalene, etc.). Physicists conjure up the three dimensional views of molecular structures while biologists describe the protein structure at four levels. The sequence of amino acids i.e., the positional information in a protein – which is the first amino acid, which is second, and so on – is called the primary structure (Figure 9.3 a) of a protein. A protein is imagined as a line, the left end represented by the first amino acid and the right end represented by the last amino.
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# 112 # BIOLOGY # (a) Primary Polypeptide # (b) Secondary Alpha – Helix Beta– plated sheet # (c) Tertiary Hydrogen Disulphide bond # (d) Quaternary Figure 9.3 Various levels of Protein Structure The first amino acid is also called as N-terminal amino acid. The last amino acid is called the C-terminal amino acid. A protein thread does not exist throughout as an extended rigid rod. The thread is folded in the form of a helix (similar to a revolving staircase). Of course, only some portions of the protein thread are arranged in the form of a helix. In proteins, only right handed helices are observed. Other regions of the protein thread are folded into other forms in what is called the secondary structure (Fig. 9.3 b). In addition, the long protein chain is also folded upon itself like a hollow woolen ball, giving rise to the tertiary structure (Fig. 9.3 c). This gives us a 3-dimensional view of a protein. Tertiary structure is absolutely necessary for the many biological activities of proteins. Some proteins are an assembly of more than one polypeptide or subunits. The manner in which these individual folded polypeptides or subunits are arranged with respect to each other (e.g. linear string of spheres, spheres arranged one upon each other in the form of a cube or plate etc.) is the architecture of a protein otherwise called the quaternary structure of a protein (Fig. 9.3 d). Adult human haemoglobin consists of 4 subunits. Two of these are identical to each other. Hence, two subunits of α type and two subunits of β type together constitute the human haemoglobin (Hb). # 9.8 ENZYMES Almost all enzymes are proteins. There are some nucleic acids that behave like enzymes. These are called ribozymes. One can depict an enzyme by a line diagram. An enzyme like any protein has a primary structure, i.e., amino acid sequence of the protein. An enzyme like any protein has the secondary and the tertiary structure. When you look at a tertiary structure (Figure 9.3 d) you will notice that the backbone of the protein chain folds.
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# BIOMOLECULES Upon itself, the chain criss-crosses itself and hence, many crevices or pockets are made. One such pocket is the ‘active site’. An active site of an enzyme is a crevice or pocket into which the substrate fits. Thus enzymes, through their active site, catalyse reactions at a high rate. Enzyme catalysts differ from inorganic catalysts in many ways, but one major difference needs mention. Inorganic catalysts work efficiently at high temperatures and high pressures, while enzymes get damaged at high temperatures (say above 40°C). However, enzymes isolated from organisms who normally live under extremely high temperatures (e.g., hot vents and sulphur springs), are stable and retain their catalytic power even at high temperatures (up to 80°-90°C). Thermal stability is thus an important quality of such enzymes isolated from thermophilic organisms. # 9.8.1 Chemical Reactions How do we understand these enzymes? Let us first understand a chemical reaction. Chemical compounds undergo two types of changes. A physical change simply refers to a change in shape without breaking of bonds. This is a physical process. Another physical process is a change in state of matter: when ice melts into water, or when water becomes a vapour. These are also physical processes. However, when bonds are broken and new bonds are formed during transformation, this will be called a chemical reaction. For example: Ba(OH)2 + HSO4 → BaSO4 + 2HO2 is an inorganic chemical reaction. Similarly, hydrolysis of starch into glucose is an organic chemical reaction. Rate of a physical or chemical process refers to the amount of product formed per unit time. It can be expressed as: rate = δP / δt Rate can also be called velocity if the direction is specified. Rates of physical and chemical processes are influenced by temperature among other factors. A general rule of thumb is that rate doubles or decreases by half for every 10°C change in either direction. Catalysed reactions proceed at rates vastly higher than that of uncatalysed ones. When enzyme catalysed reactions are observed, the rate would be vastly higher than the same but uncatalysed reaction. For example: CO2 + H2O ⟶ Carbonic anhydrase → HCO3 ← carbon dioxide water carbonic acid
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# BIOLOGY In the absence of any enzyme this reaction is very slow, with about 200 molecules of HCO3 being formed in an hour. However, by using the enzyme present within the cytoplasm called carbonic anhydrase, the reaction speeds dramatically with about 600,000 molecules being formed every second. The enzyme has accelerated the reaction rate by about 10 million times. The power of enzymes is incredible indeed! There are thousands of types of enzymes each catalysing a unique chemical or metabolic reaction. A multistep chemical reaction, when each of the steps is catalysed by the same enzyme complex or different enzymes, is called a metabolic pathway. For example, Glucose → 2 Pyruvic acid C6H12O6 + O2 → 2C3H4O3 + 2H2O is actually a metabolic pathway in which glucose becomes pyruvic acid through ten different enzyme catalysed metabolic reactions. When you study respiration in Chapter 12 you will study these reactions. At this stage you should know that this very metabolic pathway with one or two additional reactions gives rise to a variety of metabolic end products. In our skeletal muscle, under anaerobic conditions, lactic acid is formed. Under normal aerobic conditions, pyruvic acid is formed. In yeast, during fermentation, the same pathway leads to the production of ethanol (alcohol). Hence, in different conditions different products are possible. # 9.8.2 How do Enzymes bring about such High Rates of Chemical Conversions? To understand this we should study enzymes a little more. We have already understood the idea of an ‘active site’. The chemical or metabolic conversion refers to a reaction. The chemical which is converted into a product is called a ‘substrate’. Hence enzymes, i.e. proteins with three dimensional structures including an ‘active site’, convert a substrate (S) into a product (P). Symbolically, this can be depicted as: S → P It is now understood that the substrate ‘S’ has to bind the enzyme at its ‘active site’ within a given cleft or pocket. The substrate has to diffuse towards the ‘active site’. There is thus, an obligatory formation of an ‘ES’ complex. E stands for enzyme. This complex formation is a transient phenomenon. During the state where substrate is bound to the enzyme active site, a new structure of the substrate called transition state structure is formed. Very soon, after the expected bond breaking/making is completed, the product is released from the active site. In other words, the structure of substrate gets transformed into the structure of product(s). The pathway of this transformation must go through the so-called
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# BIOMOLECULES # 9.8.3 Nature of Enzyme Action Each enzyme (E) has a substrate (S) binding site in its molecule so that a highly reactive enzyme-substrate complex (ES) is produced. This complex is short-lived and dissociates into its product(s) P and the unchanged enzyme with an intermediate formation of the enzyme-product complex (EP). The formation of the ES complex is essential for catalysis. E + S ES ⍇ EP ⍇ → E + P The catalytic cycle of an enzyme action can be described in the following steps: 1. First, the substrate binds to the active site of the enzyme, fitting into the active site. 2. The binding of the substrate induces the enzyme to alter its shape, fitting more tightly around the substrate. 3. The active site of the enzyme, now in close proximity of the substrate breaks the chemical bonds of the substrate and the new enzyme-product complex is formed. Transition state structure. There could be many more ‘altered structural states’ between the stable substrate and the product. Implicit in this statement is the fact that all other intermediate structural states are unstable. Stability is something related to energy status of the molecule or the structure. Hence, when we look at this pictorially through a graph it looks like something as in Figure 9.4. The y-axis represents the potential energy content. The x-axis represents the progression of the structural transformation or states through the ‘transition state’. You would notice two things. The energy level difference between S and P. If ‘P’ is at a lower level than ‘S’, the reaction is an exothermic reaction. One need not supply energy (by heating) in order to form the product. However, whether it is an exothermic or spontaneous reaction or an endothermic or energy requiring reaction, the ‘S’ has to go through a much higher energy state or transition state. The difference in average energy content of ‘S’ from that of this transition state is called ‘activation energy’. Enzymes eventually bring down this energy barrier making the transition of ‘S’ to ‘P’ more easy.
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8124714f-1bd4-4b1c-b91a-b4a02e2c04f7
# BIOLOGY # 9.8.4 Factors Affecting Enzyme Activity The activity of an enzyme can be affected by a change in the conditions which can alter the tertiary structure of the protein. These include temperature, pH, change in substrate concentration or binding of specific chemicals that regulate its activity. # Temperature and pH Enzymes generally function in a narrow range of temperature and pH (Figure 9.5). Each enzyme shows its highest activity at a particular temperature and pH called the optimum temperature and optimum pH. Activity declines both below and above the optimum value. Low temperature preserves the enzyme in a temporarily inactive state whereas high temperature destroys enzymatic activity because proteins are denatured by heat. # Concentration of Substrate With the increase in substrate concentration, the velocity of the enzymatic reaction rises at first. The reaction ultimately reaches a maximum velocity (Vmax) which is not exceeded by any further rise in concentration of the substrate. This is because the enzyme molecules are fewer than the substrate molecules and after saturation of these molecules, there are no free enzyme molecules to bind with the additional substrate molecules (Figure 9.5). The activity of an enzyme is also sensitive to the presence of specific chemicals that bind to the enzyme. When the binding of the chemical |Enzyme activity|Velocity of reaction (V)| |---|---| |pH|Vmax| |Temperature| | |Km| | Figure 9.5 Effect of change in: (a) pH (b) Temperature and (c) Concentration of substrate on enzyme activity 2024-25
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# BIOMOLECULES shuts off enzyme activity, the process is called an inhibitor. When the inhibitor closely resembles the substrate in its molecular structure and inhibits the activity of the enzyme, it is known as competitive inhibitor. Due to its close structural similarity with the substrate, the inhibitor competes with the substrate for the substrate-binding site of the enzyme. Consequently, the substrate cannot bind and as a result, the enzyme action declines, e.g., inhibition of succinic dehydrogenase by malonate which closely resembles the substrate succinate in structure. Such competitive inhibitors are often used in the control of bacterial pathogens. # 9.8.5 Classification and Nomenclature of Enzymes Thousands of enzymes have been discovered, isolated and studied. Most of these enzymes have been classified into different groups based on the type of reactions they catalyse. Enzymes are divided into 6 classes each with 4-13 subclasses and named accordingly by a four-digit number. - Oxidoreductases/dehydrogenases: Enzymes which catalyse oxidoreduction between two substrates S and S’ e.g., S reduced + S’ oxidised ⍇ → S oxidised + S’ reduced. - Transferases: Enzymes catalysing a transfer of a group, G (other than hydrogen) between a pair of substrate S and S’ e.g., S - G + S’ ⍇ → S + S’ - G. - Hydrolases: Enzymes catalysing hydrolysis of ester, ether, peptide, glycosidic, C-C, C-halide or P-N bonds. - Lyases: Enzymes that catalyse removal of groups from substrates by mechanisms other than hydrolysis leaving double bonds. - Isomerases: Includes all enzymes catalysing inter-conversion of optical, geometric or positional isomers. - Ligases: Enzymes catalysing the linking together of 2 compounds, e.g., enzymes which catalyse joining of C-O, C-S, C-N, P-O etc. bonds. # 9.8.6 Co-factors Enzymes are composed of one or several polypeptide chains. However, there are a number of cases in which non-protein constituents called co-factors are bound to the enzyme to make the enzyme catalytically.
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# BIOLOGY active. In these instances, the protein portion of the enzymes is called the apoenzyme. Three kinds of cofactors may be identified: prosthetic groups, co-enzymes and metal ions. Prosthetic groups are organic compounds and are distinguished from other cofactors in that they are tightly bound to the apoenzyme. For example, in peroxidase and catalase, which catalyze the breakdown of hydrogen peroxide to water and oxygen, haem is the prosthetic group and it is a part of the active site of the enzyme. Co-enzymes are also organic compounds but their association with the apoenzyme is only transient, usually occurring during the course of catalysis. Furthermore, co-enzymes serve as co-factors in a number of different enzyme catalyzed reactions. The essential chemical components of many coenzymes are vitamins, e.g., coenzyme nicotinamide adenine dinucleotide (NAD) and NADP contain the vitamin niacin. A number of enzymes require metal ions for their activity which form coordination bonds with side chains at the active site and at the same time form one or more coordination bonds with the substrate, e.g., zinc is a cofactor for the proteolytic enzyme carboxypeptidase. Catalytic activity is lost when the co-factor is removed from the enzyme which testifies that they play a crucial role in the catalytic activity of the enzyme. # SUMMARY Although there is a bewildering diversity of living organisms, their chemical composition and metabolic reactions appear to be remarkably similar. The elemental composition of living tissues and non-living matter appear also to be similar when analysed qualitatively. However, a closer examination reveals that the relative abundance of carbon, hydrogen and oxygen is higher in living systems when compared to inanimate matter. The most abundant chemical in living organisms is water. There are thousands of small molecular weight (<1000 Da) biomolecules. Amino acids, monosaccharide and disaccharide sugars, fatty acids, glycerol, nucleotides, nucleosides and nitrogen bases are some of the organic compounds seen in living organisms. There are 20 types of amino acids and 5 types of nucleotides. Fats and oils are glycerides in which fatty acids are esterified to glycerol. Phospholipids contain, in addition, a phosphorylated nitrogenous compound. Only three types of macromolecules, i.e., proteins, nucleic acids and polysaccharides are found in living systems. Lipids, because of their association with membranes separate in the macromolecular fraction. Biomacromolecules are polymers. They are made of building blocks which are different. Proteins are heteropolymers made of amino acids. Nucleic acids (RNA and DNA) are composed of nucleotides. Biomacromolecules have a hierarchy of structures.
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# BIOMOLECULES primary, secondary, tertiary and quaternary. Nucleic acids serve as genetic material. Polysaccharides are components of cell wall in plants, fungi and also of the exoskeleton of arthropods. They also are storage forms of energy (e.g., starch and glycogen). Proteins serve a variety of cellular functions. Many of them are enzymes, some are antibodies, some are receptors, some are hormones and some others are structural proteins. Collagen is the most abundant protein in animal world and Ribulose bisphosphate Carboxylase-Oxygenase (RuBisCO) is the most abundant protein in the whole of the biosphere. Enzymes are proteins which catalyse biochemical reactions in the cells. Ribozymes are nucleic acids with catalytic power. Proteinaceous enzymes exhibit substrate specificity, require optimum temperature and pH for maximal activity. They are denatured at high temperatures. Enzymes lower activation energy of reactions and enhance greatly the rate of the reactions. Nucleic acids carry hereditary information and are passed on from parental generation to progeny. # EXERCISES 1. What are macromolecules? Give examples. 2. What is meant by tertiary structure of proteins? 3. Find and write down structures of 10 interesting small molecular weight biomolecules. Find if there is any industry which manufactures the compounds by isolation. Find out who are the buyers. 4. Find out and make a list of proteins used as therapeutic agents. Find other applications of proteins (e.g., Cosmetics etc.) 5. Explain the composition of triglyceride. 6. Can you attempt building models of biomolecules using commercially available atomic models (Ball and Stick models). 7. Draw the structure of the amino acid, alanine. 8. What are gums made of? Is Fevicol different? 9. Find out a qualitative test for proteins, fats and oils, amino acids and test any fruit juice, saliva, sweat and urine for them. 10. Find out how much cellulose is made by all the plants in the biosphere and compare it with how much of paper is manufactured by man and hence what is the consumption of plant material by man annually. What a loss of vegetation! 11. Describe the important properties of enzymes. 2024-25
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# UNIT 4 # PLANT PHYSIOLOGY # Chapter 11 Photosynthesis in Higher Plants # Chapter 12 Respiration in Plants # Chapter 13 Plant Growth and Development The description of structure and variation of living organisms over a period of time, ended up as two, apparently irreconcilable perspectives on biology. The two perspectives essentially rested on two levels of organisation of life forms and phenomena. One described at organismic and above level of organisation while the second described at cellular and molecular level of organisation. The first resulted in ecology and related disciplines. The second resulted in physiology and biochemistry. Description of physiological processes, in flowering plants as an example, is what is given in the chapters in this unit. The processes of photosynthesis, respiration and ultimately plant growth and development are described in molecular terms but in the context of cellular activities and even at organism level. Wherever appropriate, the relation of the physiological processes to environment is also discussed. 2024-25
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# Melvin Calvin Born in Minnesota in April, 1911, received his Ph.D. in Chemistry from the University of Minnesota. He served as Professor of Chemistry at the University of California, Berkeley. Just after World War II, when the world was under shock after the Hiroshima-Nagasaki bombings, and seeing the ill-effects of radio-activity, Calvin and co-workers put radio-activity to beneficial use. He along with J.A. Bassham studied reactions in green plants forming sugar and other substances from raw materials like carbon dioxide, water and minerals by labelling the carbon dioxide with 14C. Calvin proposed that plants change light energy to chemical energy by transferring an electron in an organised array of pigment molecules and other substances. The mapping of the pathway of carbon assimilation in photosynthesis earned him the Nobel Prize in 1961. The principles of photosynthesis as established by Calvin are, at present, being used in studies on renewable resources for energy and materials and basic studies in solar energy. Melvin Calvin research. 2024-25
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74057513-dfd3-4e64-9a6b-928351665b48
# PHOTOSYNTHESIS IN HIGHER PLANTS # 11.1 What do we Know? All animals including human beings depend on plants for their food. Have you ever wondered from where plants get their food? Green plants, in fact, have to make or rather synthesise the food they need and all other organisms depend on them for their needs. The green plants make or rather synthesise the food they need through photosynthesis and are therefore called autotrophs. # 11.2 Early Experiments You have already learnt that the autotrophic nutrition is found only in plants and all other organisms that depend on the green plants for food are heterotrophs. Green plants carry out ‘photosynthesis’, a physico-chemical process by which they use light energy to drive the synthesis of organic compounds. Ultimately, all living forms on earth depend on sunlight for energy. The use of energy from sunlight by plants doing photosynthesis is the basis of life on earth. Photosynthesis is important due to two reasons: it is the primary source of all food on earth. It is also responsible for the release of oxygen into the atmosphere by green plants. # 11.3 Where does Photosynthesis take place? What would happen if there were no oxygen to breathe? # 11.4 How many Pigments are involved in Photosynthesis? The structure of the photosynthetic machinery and the various reactions that transform light energy into chemical energy. # 11.5 What is Light Reaction? # 11.6 The Electron Transport # 11.7 Where are the ATP and NADPH Used? Let us try to find out what we already know about photosynthesis. Some simple experiments you may have done in the earlier classes have shown that chlorophyll (green pigment of the leaf), light and CO2 are required for photosynthesis to occur. # 11.8 The C4 Pathway # 11.9 Photorespiration # 11.10 Factors affecting Photosynthesis You may have carried out the experiment to look for starch in two leaves – a variegated leaf or a leaf that was partially covered with black paper, and exposed to light. On testing these leaves for the presence of starch it was clear that photosynthesis occurred only in the green parts of the leaves in the presence of light. Have you ever thought what this chapter focuses on?
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# BIOLOGY Another experiment you may have carried out where a part of a leaf is enclosed in a test tube containing some KOH soaked cotton (which absorbs CO2), while the other half is exposed to air. The setup is then placed in light for some time. On testing for the presence of starch later in the two parts of the leaf, you must have found that the exposed part of the leaf tested positive for starch while the portion that was in the tube tested negative. This showed that CO2 was required for photosynthesis. Can you explain how this conclusion could be drawn? # 11.2 EARLY EXPERIMENTS It is interesting to learn about those simple experiments that led to a gradual development in our understanding of photosynthesis. Joseph Priestley (1733-1804) in 1770 performed a series of experiments that revealed the essential role of air in the growth of green plants. Priestley, you may recall, discovered oxygen in 1774. Priestley observed that a candle burning in a closed space – a bell jar, soon gets extinguished (Figure 11.1 a, b, c, d). Similarly, a mouse would soon suffocate in a closed space. He concluded that a burning candle or an animal that breathes the air, both somehow, damage the air. But when he placed a mint plant in the same bell jar, he found that the mouse stayed alive and the candle continued to burn. Priestley hypothesised as follows: Plants restore to the air whatever breathing animals and burning candles remove. Can you imagine how Priestley would have conducted the experiment using a candle and a plant? Remember, he would need to rekindle the candle to test whether it burns after a few days. How many different ways can you think of to light the candle without disturbing the set-up? Using a similar setup as the one used by Priestley, but by placing it once in the dark and once in the sunlight, Jan Ingenhousz (1730-1799) showed that sunlight is essential to the plant process that somehow purifies the air fouled by burning candles or breathing animals. Ingenhousz in an elegant experiment with an aquatic plant showed that in bright sunlight, small bubbles were formed around the green parts while in the dark they did not. Later he identified these bubbles to be of oxygen. Hence he showed that it is only the green part of the plants that could release oxygen.
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cbda1da2-bdb3-491d-aba7-8bb2a00bb12e
# PHOTOSYNTHESIS IN HIGHER PLANTS It was not until about 1854 that Julius von Sachs provided evidence for production of glucose when plants grow. Glucose is usually stored as starch. His later studies showed that the green substance in plants (chlorophyll as we know it now) is located in special bodies (later called chloroplasts) within plant cells. He found that the green parts in plants is where glucose is made, and that the glucose is usually stored as starch. Now consider the interesting experiments done by T.W Engelmann (1843 – 1909). Using a prism he split light into its spectral components and then illuminated a green alga, *Cladophora*, placed in a suspension of aerobic bacteria. The bacteria were used to detect the sites of O2 evolution. He observed that the bacteria accumulated mainly in the region of blue and red light of the split spectrum. A first action spectrum of photosynthesis was thus described. It resembles roughly the absorption spectra of chlorophyll a and b (discussed in section 11.4). By the middle of the nineteenth century the key features of plant photosynthesis were known, namely, that plants could use light energy to make carbohydrates from CO2 and water. The empirical equation representing the total process of photosynthesis for oxygen evolving organisms was then understood as: CO2 + H2O ⟶ Light → [CH2O] + O2 where [CH2O] represented a carbohydrate (e.g., glucose, a six-carbon sugar). A milestone contribution to the understanding of photosynthesis was that made by a microbiologist, Cornelius van Niel (1897-1985), who, based on his studies of purple and green bacteria, demonstrated that photosynthesis is essentially a light-dependent reaction in which hydrogen from a suitable oxidisable compound reduces carbon dioxide to carbohydrates. This can be expressed by: 2HA + CO2 ⟶ Light → 2A + CH2O + H2O In green plants H2O is the hydrogen donor and is oxidised to O2. Some organisms do not release O2 during photosynthesis. When H2S, instead, is the hydrogen donor for purple and green sulphur bacteria, the ‘oxidation’ product is sulphur or sulphate depending on the organism and not O2. Hence, he inferred that the O2 evolved by the green plant comes from H2O, not from carbon dioxide. This was later proved by using radioisotopic techniques. The correct equation, that would represent the overall process of photosynthesis is therefore: 6CO2 + 12H2O ⟶ Light → C6H12O6 + 6H2O + 6O2 where C6H12O6 represents glucose. The O2 released is from water; this was proved using radio isotope techniques. Note that this is not a single
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# BIOLOGY reaction but description of a multistep process called photosynthesis. Can you explain why twelve molecules of water as substrate are used in the equation given above? # 11.3 WHEREDOES PHOTOSYNTHESIS TAKE PLACE? You would of course answer: in ‘the green leaf’ or ‘in the chloroplasts’, based on what you earlier read in Chapter 8. You are definitely right. Photosynthesis does take place in the green leaves of plants but it does so also in other green parts of the plants. Can you name some other parts where you think photosynthesis may occur? You would recollect from previous unit that the mesophyll cells in the leaves, have a large number of chloroplasts. Usually the chloroplasts align themselves along the walls of the mesophyll cells, such that they get the optimum quantity of the incident light. When do you think the chloroplasts will be aligned with their flat surfaces parallel to the walls? When would they be perpendicular to the incident light? You have studied the structure of chloroplast in Chapter 8. Within the chloroplast there is membranous system consisting of grana, the stroma lamellae, and the matrix stroma (Figure 11.2). There is a clear division of labour within the chloroplast. The membrane system is responsible for trapping the light energy and also for the synthesis of ATP and NADPH. In stroma, enzymatic reactions synthesise sugar, which in turn forms starch. The former set of reactions, since they are directly light driven are called light reactions (photochemical reactions). The latter are not directly light driven but are dependent on the products of light reactions (ATP and NADPH). Hence, to distinguish the latter they are called, by convention, as dark reactions (carbon reactions). However, this should not be construed to mean that they occur in darkness or that they are not light-dependent. Figure 11.2 Diagrammatic representation of an electron micrograph of a section of chloroplast 2024-25
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# PHOTOSYNTHESIS IN HIGHER PLANTS # 11.4 HOW MANY TYPES OF PIGMENTS ARE INVOLVED IN PHOTOSYNTHESIS? Looking at plants have you ever wondered why and how there are so many shades of green in their leaves – even in the same plant? We can look for an answer to this question by trying to separate the leaf pigments of any green plant through paper chromatography. A chromatographic separation of the leaf pigments shows that the colour that we see in leaves is not due to a single pigment but due to four pigments: Chlorophyll a (bright or blue green in the chromatogram), chlorophyll b (yellow green), xanthophylls (yellow) and carotenoids (yellow to yellow-orange). Let us now see what roles various pigments play in photosynthesis. Pigments are substances that have an ability to absorb light, at specific wavelengths. Can you guess which is the most abundant plant pigment in the world? Let us study the graph showing the ability of chlorophyll a pigment to absorb lights of different wavelengths (Figure 11.3 a). Of course, you are familiar with the wavelength of the visible spectrum of light as well as the VIBGYOR. From Figure 11.3a can you determine the wavelength (colour of light) at which chlorophyll a shows the maximum absorption? Does it show another absorption peak at any other wavelengths too? If yes, which one? Now look at Figure 11.3b showing the wavelengths at which maximum photosynthesis occurs in a plant. Can you see that the wavelengths at which there is maximum absorption by chlorophyll a, i.e., in the blue and the red regions, also shows higher rate of photosynthesis. Hence, we can conclude that chlorophyll a is the chief pigment associated with photosynthesis. But by looking at Figure 11.3c can you say that there is a complete one-to-one overlap between the absorption spectrum of chlorophyll a and the action spectrum of photosynthesis? # Figures - Figure 11.3a: Graph showing the absorption spectrum of chlorophyll a, b and the carotenoids - Figure 11.3b: Graph showing action spectrum of photosynthesis - Figure 11.3c: Graph showing action spectrum of photosynthesis superimposed on absorption spectrum of chlorophyll a
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# BIOLOGY These graphs, together, show that most of the photosynthesis takes place in the blue and red regions of the spectrum; some photosynthesis does take place at the other wavelengths of the visible spectrum. Let us see how this happens. Though chlorophyll is the major pigment responsible for trapping light, other thylakoid pigments like chlorophyll b, xanthophylls and carotenoids, which are called accessory pigments, also absorb light and transfer the energy to chlorophyll a. Indeed, they not only enable a wider range of wavelength of incoming light to be utilised for photosynthesis but also protect chlorophyll from photo-oxidation. # 11.5 WHAT IS LIGHT REACTION? Light reactions or the ‘Photochemical’ phase include light absorption, water splitting, oxygen release, and the formation of high-energy chemical intermediates, ATP and NADPH. Several protein complexes are involved in the process. The pigments are organised into two discrete photochemical light harvesting complexes (LHC) within the Photosystem I (PS I) and Photosystem II (PS II). These are named in the sequence of their discovery, and not in the sequence in which they function during the light reaction. The LHC are made up of hundreds of pigment molecules bound to proteins. Each photosystem has all the pigments (except one molecule of chlorophyll a) forming a light harvesting system also called antennae (Figure 11.4). These pigments help to make photosynthesis more efficient by absorbing different wavelengths of light. The single chlorophyll molecule forms the reaction centre. The reaction centre is different in both the photosystems. In PS I the reaction centre chlorophyll a has an absorption peak at 700 nm, hence is called P700, while in PS II it has absorption maxima at 680 nm, and is called P680. # 11.6 THE ELECTRON TRANSPORT In photosystem II the reaction centre chlorophyll a absorbs 680 nm wavelength of red light causing electrons to become excited and jump into an orbit farther from the atomic nucleus. These electrons are picked up by an electron acceptor which passes them to an electron transport.
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# PHOTOSYNTHESIS IN HIGHER PLANTS system consisting of cytochromes (Figure 11.5). This movement of electrons is downhill, in terms of an oxidation-reduction or redox potential scale. The electrons are not used up as they pass through the electron transport chain, but are passed on to the pigments of photosystem PS I. Simultaneously, electrons in the reaction centre of PS I are also excited when they receive red light of wavelength 700 nm and are transferred to another accepter molecule that has a greater redox potential. These electrons then are moved downhill again, this time to a molecule of energy-rich NADP+. The addition of these electrons reduces NADP+ to NADPH + H+. This whole scheme of transfer of electrons, starting from the PS II, uphill to the acceptor, down the electron transport chain to PS I, excitation of electrons, transfer to another acceptor, and finally downhill to NADP reducing it to NADPH + H+ is called the Z scheme, due to its characteristic shape (Figure 11.5). This shape is formed when all the carriers are placed in a sequence on a redox potential scale. # 11.6.1 Splitting of Water You would then ask, How does PS II supply electrons that were moved from photosystem II must be replaced. This is achieved by electrons available due to splitting of water. Water is associated with the PS II; water is split into 2H2O ⟶ 4H+ + O2 + 4e−. This creates oxygen, one of the net products of photosynthesis. The electrons needed to replace those removed from photosystem I are provided by photosystem II. We need to emphasise here that the water splitting complex is associated with the PS II, which itself is physically located on the inner side of the membrane of the thylakoid. Then, where are the protons and O2 formed likely to be released – in the lumen or on the outer side of the membrane? # 11.6.2 Cyclic and Non-cyclic Photo-phosphorylation Living organisms have the capability of extracting energy from oxidisable substances and store this in the form of bond energy. Special substances like ATP, carry this energy in their chemical bonds. The process through which
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# BIOLOGY ATP is synthesised by cells (in mitochondria and chloroplasts) is named phosphorylation. Photo-phosphorylation is the synthesis of ATP from ADP and inorganic phosphate in the presence of light. When the two photosystems work in a series, first PS II and then the PS I, a process called non-cyclic photo-phosphorylation occurs. The two photosystems are connected through an electron transport chain, as seen earlier – in the Z scheme. Both ATP and NADPH + H+ are synthesised by this kind of electron flow (Figure 11.5). When only PS I is functional, the electron is circulated within the photosystem and the phosphorylation occurs due to cyclic flow of electrons (Figure 11.6). A possible location where this could be happening is in the stroma lamellae. While the membrane or lamellae of the grana have both PS I and PS II, the stroma lamellae membranes lack PS II as well as NADP reductase enzyme. The excited electron does not pass on to NADP+ but is cycled back to the PS I complex through the electron transport chain (Figure 11.6). The cyclic flow hence results only in the synthesis of ATP, but not of NADPH + H+. Cyclic photophosphorylation also occurs when only light of wavelengths beyond 680 nm are available for excitation. # 11.6.3 Chemiosmotic Hypothesis Let us now try and understand how actually ATP is synthesised in the chloroplast. The chemiosmotic hypothesis has been put forward to explain the mechanism. Like in respiration, in photosynthesis too, ATP synthesis is linked to development of a proton gradient across a membrane. This time these are the membranes of thylakoid. There is one difference though, here the proton accumulation is towards the inside of the membrane, i.e., in the lumen. In respiration, protons accumulate in the intermembrane space of the mitochondria when electrons move through the ETS (Chapter 12). Let us understand what causes the proton gradient across the membrane. We need to consider again the processes that take place during the activation of electrons and their transport to determine the steps that cause a proton gradient to develop (Figure 11.7). (a) Since splitting of the water molecule takes place on the inner side of the membrane, the protons or hydrogen ions that are produced by the splitting of water accumulate within the lumen of the thylakoids.
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# PHOTOSYNTHESIS IN HIGHER PLANTS # Figure 11.7 ATP synthesis through chemiosmosis (b) As electrons move through the photosystems, protons are transported across the membrane. This happens because the primary accepter of electron which is located towards the outer side of the membrane transfers its electron not to an electron carrier but to an H carrier. Hence, this molecule removes a proton from the stroma while transporting an electron. When this molecule passes on its electron to the electron carrier on the inner side of the membrane, the proton is released into the inner side or the lumen side of the membrane. (c) The NADP reductase enzyme is located on the stroma side of the membrane. Along with electrons that come from the acceptor of electrons of PS I, protons are necessary for the reduction of NADP+ to NADPH + H+. These protons are also removed from the stroma. Hence, within the chloroplast, protons in the stroma decrease in number, while in the lumen there is accumulation of protons. This creates a proton gradient across the thylakoid membrane as well as a measurable decrease in pH in the lumen. Why are we so interested in the proton gradient? This gradient is important because it is the breakdown of this gradient that leads to the synthesis of ATP. The gradient is broken down due to the movement of protons across the membrane to the stroma through the transmembrane.
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# 142 # BIOLOGY channel of the CF of the ATP synthase. The ATP synthase enzyme consists of two parts: one called the CF is embedded in the thylakoid membrane and forms a transmembrane channel that carries out facilitated diffusion of protons across the membrane. The other portion is called CF and protrudes on the outer surface of the thylakoid membrane on the side that faces the stroma. The break down of the gradient provides enough energy to cause a conformational change in the CF particle of the ATP synthase, which makes the enzyme synthesise several molecules of energy-packed ATP. Chemiosmosis requires a membrane, a proton pump, a proton gradient and ATP synthase. Energy is used to pump protons across a membrane, to create a gradient or a high concentration of protons within the thylakoid lumen. ATP synthase has a channel that allows diffusion of protons back across the membrane; this releases enough energy to activate ATP synthase enzyme that catalyses the formation of ATP. Along with the NADPH produced by the movement of electrons, the ATP will be used immediately in the biosynthetic reaction taking place in the stroma, responsible for fixing CO2 and synthesis of sugars. # 11.7 WHERE ARE THE ATP AND NADPH USED? We learnt that the products of light reaction are ATP, NADPH and O2. Of these O2 diffuses out of the chloroplast while ATP and NADPH are used to drive the processes leading to the synthesis of food, more accurately, sugars. This is the biosynthetic phase of photosynthesis. This process does not directly depend on the presence of light but is dependent on the products of the light reaction, i.e., ATP and NADPH, besides CO2 and H2O. You may wonder how this could be verified; it is simple: immediately after light becomes unavailable, the biosynthetic process continues for some time, and then stops. If then, light is made available, the synthesis starts again. Can we, hence, say that calling the biosynthetic phase as the dark reaction is a misnomer? Discuss this amongst yourselves. Let us now see how the ATP and NADPH are used in the biosynthetic phase. We saw earlier that CO2 is combined with H2O to produce (CHO)n or sugars. It was of interest to scientists to find out how this reaction proceeded, or rather what was the first product formed when CO2 is taken into a reaction or fixed. Just after world war II, among the several efforts to put radioisotopes to beneficial use, the work of Melvin Calvin is exemplary. The use of radioactive 14C by him in algal photosynthesis studies led to the discovery that the first CO2 fixation product was a 3-carbon organic acid. He also contributed to working out the complete biosynthetic pathway; hence it was called Calvin cycle after him. The first product identified was 3-phosphoglyceric acid or in short PGA. How many carbon atoms does it have? 2024-25
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# PHOTOSYNTHESIS IN HIGHER PLANTS Scientists also tried to know whether all plants have PGA as the first product of CO2 fixation, or whether any other product was formed in other plants. Experiments conducted over a wide range of plants led to the discovery of another group of plants, where the first stable product of CO2 fixation was again an organic acid, but one which had 4 carbon atoms in it. This acid was identified to be oxaloacetic acid or OAA. Since then CO assimilation during photosynthesis was said to be of two main types: those plants in which the first product of CO2 fixation is a C3 acid (PGA), i.e., the C3 pathway, and those in which the first product was a C4 acid (OAA), i.e., the C4 pathway. These two groups of plants showed other associated characteristics that we will discuss later. # 11.7.1 The Primary Acceptor of CO2 Let us now ask ourselves a question that was asked by the scientists who were struggling to understand the ‘dark reaction’. How many carbon atoms would a molecule have which after accepting (fixing) CO2, would have 3 carbons (of PGA)? The studies very unexpectedly showed that the acceptor molecule was a 5-carbon ketose sugar – ribulose bisphosphate (RuBP). Did any of you think of this possibility? Do not worry; the scientists also took a long time and conducted many experiments to reach this conclusion. They also believed that since the first product was a C3 acid, the primary acceptor would be a 2-carbon compound; they spent many years trying to identify a 2-carbon compound before they discovered the 5-carbon RuBP. # 11.7.2 The Calvin Cycle Calvin and his co-workers then worked out the whole pathway and showed that the pathway operated in a cyclic manner; the RuBP was regenerated. Let us now see how the Calvin pathway operates and where the sugar is synthesised. Let us at the outset understand very clearly that the Calvin pathway occurs in all photosynthetic plants; it does not matter whether they have C3 or C4 (or any other) pathways. For ease of understanding, the Calvin cycle can be described under three stages: carboxylation, reduction and regeneration. 1. Carboxylation – Carboxylation is the fixation of CO2 into a stable organic intermediate. Carboxylation is the most crucial step of the Calvin cycle where CO2 is utilised for the carboxylation of RuBP. This reaction is catalysed by the enzyme RuBP carboxylase which results in the formation of two molecules of 3-PGA. Since this enzyme also has an oxygenation activity it would be more correct to call it RuBP carboxylase-oxygenase or RuBisCO.
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# BIOLOGY # Atmosphere |Ribulose-1,5-bisphosphate|CO2 + H2O| |---|---| |Carboxylation|ADP| |Regeneration|3-phosphoglycerate| |ATP|ATP| |Reduction|NADPH| |Triose phosphate|ADP + P + NADP+| |Sucrose, starch|Sucrose, starch| Figure 11.8 The Calvin cycle proceeds in three stages: (1) carboxylation, during which CO2 combines with ribulose-1,5-bisphosphate; (2) reduction, during which carbohydrate is formed at the expense of the photochemically made ATP and NADPH; and (3) regeneration during which the CO2 acceptor ribulose-1,5-bisphosphate is formed again so that the cycle continues. 1. Reduction – These are a series of reactions that lead to the formation of glucose. The steps involve utilisation of 2 molecules of ATP for phosphorylation and two of NADPH for reduction per CO2 molecule fixed. The fixation of six molecules of CO2 and 6 turns of the cycle are required for the formation of one molecule of glucose from the pathway. 2. Regeneration – Regeneration of the CO2 acceptor molecule RuBP is crucial if the cycle is to continue uninterrupted. The regeneration steps require one ATP for phosphorylation to form RuBP. 2024-25
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# PHOTOSYNTHESIS IN HIGHER PLANTS Hence for every CO2 molecule entering the Calvin cycle, 3 molecules of ATP and 2 of NADPH are required. It is probably to meet this difference in number of ATP and NADPH used in the dark reaction that the cyclic phosphorylation takes place. To make one molecule of glucose 6 turns of the cycle are required. Work out how many ATP and NADPH molecules will be required to make one molecule of glucose through the Calvin pathway. It might help you to understand all of this if we look at what goes in and what comes out of the Calvin cycle. |In|Out| |---|---| |Six CO2|One glucose| |18 ATP|18 ADP| |12 NADPH|12 NADP| # 11.8 THE C4 PATHWAY Plants that are adapted to dry tropical regions have the C4 pathway mentioned earlier. Though these plants have the C4 oxaloacetic acid as the first CO2 fixation product they use the C3 pathway or the Calvin cycle as the main biosynthetic pathway. Then, in what way are they different from C3 plants? This is a question that you may reasonably ask. C4 plants are special: They have a special type of leaf anatomy, they tolerate higher temperatures, they show a response to high light intensities, they lack a process called photorespiration and have greater productivity of biomass. Let us understand these one by one. Study vertical sections of leaves, one of a C3 plant and the other of a C4 plant. Do you notice the differences? Do both have the same types of mesophylls? Do they have similar cells around the vascular bundle sheath? The particularly large cells around the vascular bundles of the C4 plants are called bundle sheath cells, and the leaves which have such anatomy are said to have ‘Kranz’ anatomy. ‘Kranz’ means ‘wreath’ and is a reflection of the arrangement of cells. The bundle sheath cells may form several layers around the vascular bundles; they are characterised by having a large number of chloroplasts, thick walls impervious to gaseous exchange and no intercellular spaces. You may like to cut a section of the leaves of C4 plants – maize or sorghum – to observe the Kranz anatomy and the distribution of mesophyll cells. It would be interesting for you to collect leaves of diverse species of plants around you and cut vertical sections of the leaves. Observe under the microscope – look for the bundle sheath around the vascular bundles. The presence of the bundle sheath would help you identify the C4 plants.
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# BIOLOGY Now study the pathway shown in Figure 11.9. This pathway that has been named the Hatch and Slack Pathway, is again a cyclic process. Let us study the pathway by listing the steps. The primary CO2 acceptor is a 3-carbon molecule phosphoenol pyruvate (PEP) and is present in the mesophyll cells. The enzyme responsible for this fixation is PEP carboxylase or PEPcase. It is important to register that the mesophyll cells lack RuBisCO enzyme. The C4 acid OAA is formed in the mesophyll cells. It then forms other 4-carbon compounds like malic acid or aspartic acid in the mesophyll cells itself, which are transported to the bundle sheath cells. In the bundle sheath cells these C acids are broken down to release CO2 and a 3-carbon molecule. The 3-carbon molecule is transported back to the mesophyll where it is converted to PEP again, thus, completing the cycle. The CO2 released in the bundle sheath cells enters the C3 pathway, a pathway common to all plants. Figure 11.9 Diagrammatic representation of the Hatch and Slack Pathway
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# PHOTOSYNTHESIS IN HIGHER PLANTS rich in an enzyme Ribulose bisphosphate carboxylase-oxygenase (RuBisCO), but lack PEPcase. Thus, the basic pathway that results in the formation of the sugars, the Calvin pathway, is common to the C3 and C4 plants. Did you note that the Calvin pathway occurs in all the mesophyll cells of the C3 plants? In the C4 plants it does not take place in the mesophyll cells but does so only in the bundle sheath cells. # 11.9 PHOTORESPIRATION Let us try and understand one more process that creates an important difference between C3 and C4 plants – Photorespiration. To understand photorespiration we have to know a little bit more about the first step of the Calvin pathway – the first CO2 fixation step. This is the reaction where RuBP combines with CO2 to form 2 molecules of 3PGA, that is catalysed by RuBisCO. RuBP + CO2 ⟶ 2 3PGA RuBisCO that is the most abundant enzyme in the world (Do you wonder why?) is characterised by the fact that its active site can bind to both CO2 and O2 – hence the name. Can you think how this could be possible? RuBisCO has a much greater affinity for CO2 when the CO2: O2 ratio is nearly equal. Imagine what would happen if this were not so! This binding is competitive. It is the relative concentration of O2 and CO2 that determines which of the two will bind to the enzyme. In C3 plants some O2 does bind to RuBisCO, and hence CO2 fixation is decreased. Here the RuBP instead of being converted to 2 molecules of PGA binds with O2 to form one molecule of phosphoglycerate and phosphoglycolate (2 Carbon) in a pathway called photorespiration. In the photorespiratory pathway, there is neither synthesis of sugars, nor of ATP. Rather it results in the release of CO2 with the utilisation of ATP. In the photorespiratory pathway there is no synthesis of ATP or NADPH. The biological function of photorespiration is not known yet. In C4 plants photorespiration does not occur. This is because they have a mechanism that increases the concentration of CO2 at the enzyme site. This takes place when the C4 acid from the mesophyll is broken down in the bundle sheath cells to release CO2 – this results in increasing the intracellular concentration of CO2. In turn, this ensures that the RuBisCO functions as a carboxylase minimising the oxygenase activity. Now that you know that the C4 plants lack photorespiration, you probably can understand why productivity and yields are better in these plants. In addition these plants show tolerance to higher temperatures. Based on the above discussion can you compare plants showing the C3 and the C4 pathway? Use the table format given in table 11.1 and fill in the information.
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# BIOLOGY # TABLE 11.1 Fill in the Columns 2 and 3 in this table to highlight the differences between C and C4 Plants |Characteristics|C3 Plants|C4 Plants|Choose from| |---|---|---|---| |Cell type in which the Calvin cycle takes place| | |Mesophyll/Bundle sheath/both| |Cell type in which the initial carboxylation reaction occurs| | |Mesophyll/Bundle sheath/both| |How many cell types does the leaf have that fix CO2|Two: Bundle sheath and mesophyll|One: Mesophyll|Three: Bundle sheath, palisade, spongy mesophyll| |Which is the primary CO2 acceptor|RuBP|PEP/PGA| | |Number of carbons in the primary CO2 acceptor|5 / 4 / 3| | | |Which is the primary CO2 fixation product|PGA|OAA/RuBP/PEP| | |No. of carbons in the primary CO fixation product|3 / 4 / 5| | | |Does the plant have RuBisCO?|Yes|No/Not always| | |Does the plant have PEP Case?|Yes|No/Not always| | |Which cells in the plant have Rubisco?|Mesophyll|Bundle sheath/none| | |CO fixation rate under high light conditions| |Low/high/medium| | |Whether photorespiration is present at low light intensities| |High/negligible/sometimes| | |Whether photorespiration is present at high light intensities| |High/negligible/sometimes| | |Whether photorespiration would be present at low CO2 concentrations| |High/negligible/sometimes| | |Whether photorespiration would be present at high CO2 concentrations| |High/negligible/sometimes| | |Temperature optimum|30-40 C|20-25 C/above 40 C| | |Examples|Cut vertical sections of leaves of different plants and observe under the microscope for Kranz anatomy and list them in the appropriate columns.|Cut vertical sections of leaves of different plants and observe under the microscope for Kranz anatomy and list them in the appropriate columns.|Cut vertical sections of leaves of different plants and observe under the microscope for Kranz anatomy and list them in the appropriate columns.|
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# PHOTOSYNTHESIS IN HIGHER PLANTS # 11.10 FACTORS AFFECTING PHOTOSYNTHESIS An understanding of the factors that affect photosynthesis is necessary. The rate of photosynthesis is very important in determining the yield of plants including crop plants. Photosynthesis is under the influence of several factors, both internal (plant) and external. The plant factors include the number, size, age and orientation of leaves, mesophyll cells and chloroplasts, internal CO2 concentration and the amount of chlorophyll. The plant or internal factors are dependent on the genetic predisposition and the growth of the plant. The external factors would include the availability of sunlight, temperature, CO2 concentration and water. As a plant photosynthesises, all these factors will simultaneously affect its rate. Hence, though several factors interact and simultaneously affect photosynthesis or CO2 fixation, usually one factor is the major cause or is the one that limits the rate. Hence, at any point the rate will be determined by the factor available at sub-optimal levels. When several factors affect any [bio] chemical process, Blackman’s (1905) Law of Limiting Factors comes into effect. This states the following: If a chemical process is affected by more than one factor, then its rate will be determined by the factor which is nearest to its minimal value: it is the factor which directly affects the process if its quantity is changed. For example, despite the presence of a green leaf and optimal light and CO2 conditions, the plant may not photosynthesise if the temperature is very low. This leaf, if given the optimal temperature, will start photosynthesising. # 11.10.1 Light We need to distinguish between light quality, light intensity and the duration of exposure to light, while discussing light as a factor that affects photosynthesis. There is a linear relationship between incident light and CO2 fixation rates at low light intensities. At higher light intensities, gradually the rate does not show further increase as other factors become limiting (Figure 11.10). What is interesting to note is that light saturation occurs at 10 per cent of the full sunlight. Hence, except for plants in shade or in dense forests, light is rarely a limiting factor in nature. Increase in rate of photosynthesis. |A|B|C|D|E| |---|---|---|---|---| |Light intensity|Light intensity|Light intensity|Light intensity|Light intensity| Figure 11.10 Graph of light intensity on the rate of photosynthesis
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# 150 # BIOLOGY Incident light beyond a point causes the breakdown of chlorophyll and a decrease in photosynthesis. # 11.10.2 Carbon dioxide Concentration Carbon dioxide is the major limiting factor for photosynthesis. The concentration of CO2 is very low in the atmosphere (between 0.03 and 0.04 per cent). Increase in concentration up to 0.05 per cent can cause an increase in CO2 fixation rates; beyond this the levels can become damaging over longer periods. The C3 and C4 plants respond differently to CO2 concentrations. At low light conditions neither group responds to high CO2 conditions. At high light intensities, both C3 and C4 plants show increase in the rates of photosynthesis. What is important to note is that the C3 plants show saturation at about 360 μlL-1 while C4 responds to increased CO2 concentration and saturation is seen only beyond 450 μlL-1. Thus, current availability of CO2 levels is limiting to the C4 plants. The fact that C4 plants respond to higher CO2 concentration by showing increased rates of photosynthesis leading to higher productivity has been used for some greenhouse crops such as tomatoes and bell pepper. They are allowed to grow in carbon dioxide enriched atmosphere that leads to higher yields. # 11.10.3 Temperature The dark reactions being enzymatic are temperature controlled. Though the light reactions are also temperature sensitive they are affected to a much lesser extent. The C4 plants respond to higher temperatures and show higher rate of photosynthesis while C3 plants have a much lower temperature optimum. The temperature optimum for photosynthesis of different plants also depends on the habitat that they are adapted to. Tropical plants have a higher temperature optimum than the plants adapted to temperate climates. # 11.10.4 Water Even though water is one of the reactants in the light reaction, the effect of water as a factor is more through its effect on the plant, rather than directly on photosynthesis. Water stress causes the stomata to close hence reducing the CO2 availability. Besides, water stress also makes leaves wilt, thus, reducing the surface area of the leaves and their metabolic activity as well. 2024-25
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# PHOTOSYNTHESIS IN HIGHER PLANTS # SUMMARY Green plants make their own food by photosynthesis. During this process carbon dioxide from the atmosphere is taken in by leaves through stomata and used for making carbohydrates, principally glucose and starch. Photosynthesis takes place only in the green parts of the plants, mainly the leaves. Within the leaves, the mesophyll cells have a large number of chloroplasts that are responsible for CO2 fixation. Within the chloroplasts, the membranes are sites for the light reaction, while the chemosynthetic pathway occurs in the stroma. Photosynthesis has two stages: the light reaction and the carbon fixing reactions. In the light reaction the light energy is absorbed by the pigments present in the antenna, and funnelled to special chlorophyll a molecules called reaction centre chlorophylls. There are two photosystems, PS I and PS II. PS I has a 700 nm absorbing chlorophyll a P700 molecule at its reaction centre, while PS II has a P680 reaction centre that absorbs red light at 680 nm. After absorbing light, electrons are excited and transferred through PS II and PS I and finally to NAD forming NADH. During this process a proton gradient is created across the membrane of the thylakoid. The breakdown of the protons gradient due to movement through the F0 part of the ATPase enzyme releases enough energy for synthesis of ATP. Splitting of water molecules is associated with PS II resulting in the release of O2, protons and transfer of electrons to PS II. In the carbon fixation cycle, CO2 is added by the enzyme, RuBisCO, to a 5-carbon compound RuBP that is converted to 2 molecules of 3-carbon PGA. This is then converted to sugar by the Calvin cycle, and the RuBP is regenerated. During this process ATP and NADPH synthesised in the light reaction are utilised. RuBisCO also catalyses a wasteful oxygenation reaction in C3 plants: photorespiration. Some tropical plants show a special type of photosynthesis called C4 pathway. In these plants the first product of CO2 fixation that takes place in the mesophyll, is a 4-carbon compound. In the bundle sheath cells the Calvin pathway is carried out for the synthesis of carbohydrates. # EXERCISES 1. By looking at a plant externally can you tell whether a plant is C3 or C4? Why and how? 2. By looking at which internal structure of a plant can you tell whether a plant is C3 or C4? Explain. 3. Even though a very few cells in a C4 plant carry out the biosynthetic – Calvin pathway, yet they are highly productive. Can you discuss why? 2024-25
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# BIOLOGY 1. RuBisCO is an enzyme that acts both as a carboxylase and oxygenase. Why do you think RuBisCO carries out more carboxylation in C3 plants? 2. Suppose there were plants that had a high concentration of Chlorophyll b, but lacked chlorophyll a, would it carry out photosynthesis? Then why do plants have chlorophyll b and other accessory pigments? 3. Why is the colour of a leaf kept in the dark frequently yellow, or pale green? Which pigment do you think is more stable? 4. Look at leaves of the same plant on the shady side and compare it with the leaves on the sunny side. Or, compare the potted plants kept in the sunlight with those in the shade. Which of them has leaves that are darker green? Why? 5. Figure 11.10 shows the effect of light on the rate of photosynthesis. Based on the graph, answer the following questions: 1. (a) At which point/s (A, B or C) in the curve is light a limiting factor? 2. (b) What could be the limiting factor/s in region A? 3. (c) What do C and D represent on the curve? 6. Give comparison between the following: 1. (a) C3 and C4 pathways 2. (b) Cyclic and non-cyclic photophosphorylation 3. (c) Anatomy of leaf in C3 and C4 plants 2024-25
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# CHAPTER 10 # CELL CYCLE AND CELL DIVISION # 10.1 Cell Cycle Are you aware that all organisms, even the largest, start their life from a single cell? You may wonder how a single cell then goes on to form such large organisms. Growth and reproduction are characteristics of cells, indeed of all living organisms. All cells reproduce by dividing into two, with each parental cell giving rise to two daughter cells each time they divide. These newly formed daughter cells can themselves grow and divide, giving rise to a new cell population that is formed by the growth and division of a single parental cell and its progeny. In other words, such cycles of growth and division allow a single cell to form a structure consisting of millions of cells. # 10.2 M Phase Cell division is a very important process in all living organisms. During the division of a cell, DNA replication and cell growth also take place. All these processes, i.e., cell division, DNA replication, and cell growth, hence, have to take place in a coordinated way to ensure correct division and formation of progeny cells containing intact genomes. The sequence of events by which a cell duplicates its genome, synthesises the other constituents of the cell and eventually divides into two daughter cells is termed cell cycle. Although cell growth (in terms of cytoplasmic increase) is a continuous process, DNA synthesis occurs only during one specific stage in the cell cycle. The replicated chromosomes (DNA) are then distributed to daughter nuclei by a complex series of events during cell division. These events are themselves under genetic control. # 10.3 Significance of Mitosis 2024-25
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# CELL CYCLE AND CELL DIVISION # 10.1.1 Phases of Cell Cycle A typical eukaryotic cell cycle is illustrated by human cells in culture. These cells divide once in approximately every 24 hours (Figure 10.1). However, this duration of cell cycle can vary from organism to organism and also from cell type to cell type. Yeast for example, can progress through the cell cycle in only about 90 minutes. The cell cycle is divided into two basic phases: - Interphase - M Phase # M Phase (Mitosis phase) The M Phase represents the phase when the actual cell division or mitosis occurs and the interphase represents the phase between two successive M phases. It is significant to note that in the 24 hour average duration of cell cycle of a human cell, cell division proper lasts for only about an hour. The interphase lasts more than 95% of the duration of cell cycle. The M Phase starts with the nuclear division, corresponding to the separation of daughter chromosomes (karyokinesis) and usually ends with division of cytoplasm (cytokinesis). The interphase, though called the resting phase, is the time during which the cell is preparing for division by undergoing both cell growth and DNA replication in an orderly manner. The interphase is divided into three further phases: - G1 phase (Gap 1) - S phase (Synthesis) - G2 phase (Gap 2) G1 phase corresponds to the interval between mitosis and initiation of DNA replication. During G1 phase the cell is metabolically active and continuously grows but does not replicate its DNA. S or synthesis phase marks the period during which DNA synthesis or replication takes place. During this time the amount of DNA per cell doubles. If the initial amount of DNA is denoted as 2C then it increases to 4C. However, there is no increase in the chromosome number; if the cell had diploid or 2n number of chromosomes at G1, even after S phase the number of chromosomes remains the same, i.e., 2n. In animal cells, during the S phase, DNA replication begins in the nucleus, and the centriole duplicates in the cytoplasm. During the G2 phase, proteins are synthesised in preparation for mitosis while cell growth continues. # How do plants and animals continue to grow all their lives? Do all cells in a plant divide all the time? Do you think all cells continue to divide in all plants and animals? Can you tell the name and the location of tissues having cells that divide all their life in higher plants? Do animals have similar meristematic tissues?
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c223ad2c-54be-414c-82b5-a2633df40d8e
# BIOLOGY You have studied mitosis in onion root tip cells. It has 16 chromosomes in each cell. Can you tell how many chromosomes will the cell have at G1 phase, after S phase, and after M phase? Also, what will be the DNA content of the cells at G1, if the content after M phase is 2C? Some cells in the adult animals do not appear to exhibit division (e.g., heart cells) and many other cells divide only occasionally, as needed to replace cells that have been lost because of injury or cell death. These cells that do not divide further exit G phase to enter an inactive stage called quiescent stage (G0) of the cell cycle. Cells in this stage remain metabolically active but no longer proliferate unless called on to do so depending on the requirement of the organism. In animals, mitotic cell division is only seen in the diploid somatic cells. However, there are few exceptions to this where haploid cells divide by mitosis, for example, male honey bees. Against this, the plants can show mitotic divisions in both haploid and diploid cells. From your recollection of examples of alternation of generations in plants (Chapter 3) identify plant species and stages at which mitosis is seen in haploid cells. # 10.2 M PHASE This is the most dramatic period of the cell cycle, involving a major reorganisation of virtually all components of the cell. Since the number of chromosomes in the parent and progeny cells is the same, it is also called as equational division. Though for convenience mitosis has been divided into four stages of nuclear division (karyokinesis), it is very essential to understand that cell division is a progressive process and very clear-cut lines cannot be drawn between various stages. Karyokinesis involves following four stages: # Prophase # Metaphase # Anaphase # Telophase # 10.2.1 Prophase Prophase which is the first stage of karyokinesis of mitosis follows the S and G phases of interphase. In the S and G2 phases the new DNA molecules formed are not distinct but intertwined. Prophase is marked by the initiation of condensation of chromosomal material. The chromosomal material becomes untangled during the process of chromatin condensation. The centrosome, which had undergone duplication during S phase of interphase, now begins to move towards opposite poles of the cell. The completion of prophase can thus be marked by the following characteristic events: - Chromosomal material condenses to form compact mitotic chromosomes. Chromosomes are seen to be composed of two chromatids attached together at the centromere. - Centrosome which had undergone duplication during interphase, begins to move towards opposite poles of the cell. Each centrosome radiates out microtubules called asters. The two asters together with spindle fibres forms mitotic apparatus.
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b811ad80-ec01-44c4-a895-e9b235666142
# CELL CYCLE AND CELL DIVISION Cells at the end of prophase, when viewed under the microscope, do not show golgi complexes, endoplasmic reticulum, nucleolus and the nuclear envelope. # 10.2.2 Metaphase The complete disintegration of the nuclear envelope marks the start of the second phase of mitosis, hence the chromosomes are spread through the cytoplasm of the cell. By this stage, condensation of chromosomes is completed and they can be observed clearly under the microscope. This then, is the stage at which morphology of chromosomes is most easily studied. At this stage, metaphase chromosome is made up of two sister chromatids, which are held together by the centromere (Figure 10.2 b). Small disc-shaped structures at the surface of the centromeres are called kinetochores. These structures serve as the sites of attachment of spindle fibres (formed by the spindle fibres) to the chromosomes that are moved into position at the centre of the cell. Hence, the metaphase is characterised by all the chromosomes coming to lie at the equator with one chromatid of each chromosome connected by its kinetochore to spindle fibres from one pole and its sister chromatid connected by its kinetochore to spindle fibres from the opposite pole (Figure 10.2 b). The plane of alignment of the chromosomes at metaphase is referred to as the metaphase plate. The key features of metaphase are: - Spindle fibres attach to kinetochores of chromosomes. - Chromosomes are moved to spindle equator and get aligned along metaphase plate through spindle fibres to both poles. # 10.2.3 Anaphase At the onset of anaphase, each chromosome arranged at the metaphase plate is split simultaneously and the two daughter chromatids, now referred to as daughter chromosomes of the future daughter nuclei, begin their migration towards the two opposite poles. As each chromosome moves away from the equatorial plate, the centromere of each chromosome remains directed towards the pole and hence at the leading edge, with the arms of the chromosome trailing behind (Figure 10.2 c). Thus, anaphase stage is characterised by Figure 10.2 a and b: A diagrammatic view of stages in mitosis
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d348ce19-210b-4669-b476-6c8332da17ff
# BIOLOGY # 10.2.4 Telophase At the beginning of the final stage of karyokinesis, i.e., telophase, the chromosomes that have reached their respective poles decondense and lose their individuality. The individual chromosomes can no longer be seen and each set of chromatin material tends to collect at each of the two poles (Figure 10.2 d). This is the stage which shows the following key events: - Chromosomes cluster at opposite spindle poles and their identity is lost as discrete elements. - Nuclear envelope develops around the chromosome clusters at each pole forming two daughter nuclei. - Nucleolus, golgi complex and ER reform. # 10.2.5 Cytokinesis Mitosis accomplishes not only the segregation of duplicated chromosomes into daughter nuclei (karyokinesis), but the cell itself is divided into two daughter cells by the separation of cytoplasm called cytokinesis at the end of which cell division gets completed (Figure 10.2 e). In an animal cell, this is achieved by the appearance of a furrow in the plasma membrane. The furrow gradually deepens and ultimately joins in the centre dividing the cell cytoplasm into two. Plant cells however, are enclosed by a relatively inextensible cell wall, therefore they undergo cytokinesis by a different mechanism. In plant cells, wall formation starts in the centre of the cell and grows outward to meet the existing lateral walls. The formation of the new cell wall begins with the formation of a simple precursor, called the cell-plate that represents the middle lamella between the walls of two adjacent cells. At the time of cytoplasmic division, organelles like mitochondria and plastids get distributed between the two daughter cells. In some organisms karyokinesis is not followed by cytokinesis as a result of which multinucleate condition arises leading to the formation of syncytium (e.g., liquid endosperm in coconut).
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564c8a73-1fce-4d63-bdbb-18616561191c
# CELL CYCLE AND CELL DIVISION # 10.3 Significance of Mitosis Mitosis or the equational division is usually restricted to the diploid cells only. However, in some lower plants and in some social insects haploid cells also divide by mitosis. It is very essential to understand the significance of this division in the life of an organism. Are you aware of some examples where you have studied about haploid and diploid insects? Mitosis usually results in the production of diploid daughter cells with identical genetic complement. The growth of multicellular organisms is due to mitosis. Cell growth results in disturbing the ratio between the nucleus and the cytoplasm. It therefore becomes essential for the cell to divide to restore the nucleo-cytoplasmic ratio. A very significant contribution of mitosis is cell repair. The cells of the upper layer of the epidermis, cells of the lining of the gut, and blood cells are being constantly replaced. Mitotic divisions in the meristematic tissues – the apical and the lateral cambium, result in a continuous growth of plants throughout their life. # 10.4 MEIOSIS The production of offspring by sexual reproduction includes the fusion of two gametes, each with a complete haploid set of chromosomes. Gametes are formed from specialised diploid cells. This specialised kind of cell division that reduces the chromosome number by half results in the production of haploid daughter cells. This kind of division is called meiosis. Meiosis ensures the production of haploid phase in the life cycle of sexually reproducing organisms whereas fertilisation restores the diploid phase. We come across meiosis during gametogenesis in plants and animals. This leads to the formation of haploid gametes. The key features of meiosis are as follows: - Meiosis involves two sequential cycles of nuclear and cell division called meiosis I and meiosis II but only a single cycle of DNA replication. - Meiosis I is initiated after the parental chromosomes have replicated to produce identical sister chromatids at the S phase. - Meiosis involves pairing of homologous chromosomes and recombination between non-sister chromatids of homologous chromosomes. - Four haploid cells are formed at the end of meiosis II. Meiotic events can be grouped under the following phases: |Meiosis I|Meiosis II| |---|---| |Prophase I|Prophase II| |Metaphase I|Metaphase II| |Anaphase I|Anaphase II| |Telophase I|Telophase II|
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da7dfce8-34e2-4a06-8ed6-edba1e5daca7
# BIOLOGY # 10.4.1 Meiosis I Prophase I: Prophase of the first meiotic division is typically longer and more complex when compared to prophase of mitosis. It has been further subdivided into the following five phases based on chromosomal behaviour, i.e., Leptotene, Zygotene, Pachytene, Diplotene and Diakinesis. During leptotene stage the chromosomes become gradually visible under the light microscope. The compaction of chromosomes continues throughout leptotene. This is followed by the second stage of prophase I called zygotene. During this stage chromosomes start pairing together and this process of association is called synapsis. Such paired chromosomes are called homologous chromosomes. Electron micrographs of this stage indicate that chromosome synapsis is accompanied by the formation of complex structure called synaptonemal complex. The complex formed by a pair of synapsed homologous chromosomes is called a bivalent or a tetrad. However, these are more clearly visible at the next stage. The first two stages of prophase I are relatively short-lived compared to the next stage that is pachytene. During this stage, the four chromatids of each bivalent chromosomes becomes distinct and clearly appears as tetrads. This stage is characterised by the appearance of recombination nodules, the sites at which crossing over occurs between non-sister chromatids of the homologous chromosomes. Crossing over is the exchange of genetic material between two homologous chromosomes. Crossing over is also an enzyme-mediated process and the enzyme involved is called recombinase. Crossing over leads to recombination of genetic material on the two chromosomes. Recombination between homologous chromosomes is completed by the end of pachytene, leaving the chromosomes linked at the sites of crossing over. The beginning of diplotene is recognised by the dissolution of the synaptonemal complex and the tendency of the recombined homologous chromosomes of the bivalents to separate from each other except at the sites of crossovers. These X-shaped structures, are called chiasmata. In oocytes of some vertebrates, diplotene can last for months or years. The final stage of meiotic prophase I is diakinesis. This is marked by terminalisation of chiasmata. During this phase the chromosomes are fully condensed and the meiotic spindle is assembled to prepare the homologous chromosomes for separation. By the end of diakinesis, the nucleolus disappears and the nuclear envelope also breaks down. Diakinesis represents transition to metaphase. Metaphase I: The bivalent chromosomes align on the equatorial plate (Figure 10.3). The microtubules from the opposite poles of the spindle attach to the kinetochore of homologous chromosomes.
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# CELL CYCLE AND CELL DIVISION # 10.4.1 Meiosis I Anaphase I: The homologous chromosomes separate, while sister chromatids remain associated at their centromeres (Figure 10.3). Telophase I: The nuclear membrane and nucleolus reappear, cytokinesis follows and this is called as dyad of cells (Figure 10.3). Although in many cases the chromosomes do undergo some dispersion, they do not reach the extremely extended state of the interphase nucleus. The stage between the two meiotic divisions is called interkinesis and is generally short lived. There is no replication of DNA during interkinesis. Interkinesis is followed by prophase II, a much simpler prophase than prophase I. # 10.4.2 Meiosis II Prophase II: Meiosis II is initiated immediately after cytokinesis, usually before the chromosomes have fully elongated. In contrast to meiosis I, meiosis II resembles a normal mitosis. The nuclear membrane disappears by the end of prophase II (Figure 10.4). The chromosomes again become compact. Metaphase II: At this stage the chromosomes align at the equator and the microtubules from opposite poles of the spindle get attached to the kinetochores (Figure 10.4) of sister chromatids. Anaphase II: It begins with the simultaneous splitting of the centromere of each chromosome (which was holding the sister chromatids together), allowing them to move toward opposite poles of the cell (Figure 10.4) by shortening of microtubules attached to kinetochores. 2024-25
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# 128 # BIOLOGY # Figure 10.4 Stages of Meiosis II Telophase II: Meiosis ends with telophase II, in which the two groups of chromosomes once again get enclosed by a nuclear envelope; cytokinesis follows resulting in the formation of tetrad of cells i.e., four haploid daughter cells (Figure 10.4). # 10.5 SIGNIFICANCE OF MEIOSIS Meiosis is the mechanism by which conservation of specific chromosome number of each species is achieved across generations in sexually reproducing organisms, even though the process, per se, paradoxically, results in reduction of chromosome number by half. It also increases the genetic variability in the population of organisms from one generation to the next. Variations are very important for the process of evolution. # SUMMARY According to the cell theory, cells arise from preexisting cells. The process by which this occurs is called cell division. Any sexually reproducing organism starts its life cycle from a single-celled zygote. Cell division does not stop with the formation of the mature organism but continues throughout its life cycle. 2024-25
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36026b84-90f1-4aaa-8337-13af93909a07
# CELL CYCLE AND CELL DIVISION The stages through which a cell passes from one division to the next is called the cell cycle. Cell cycle is divided into two phases called (i) Interphase – a period of preparation for cell division, and (ii) Mitosis (M phase) – the actual period of cell division. Interphase is further subdivided into G1, S and G2. G1 phase is the period when the cell grows and carries out normal metabolism. Most of the organelle duplication also occurs during this phase. S phase marks the phase of DNA replication and chromosome duplication. G2 phase is the period of cytoplasmic growth. Mitosis is also divided into four stages namely prophase, metaphase, anaphase and telophase. Chromosome condensation occurs during prophase. Simultaneously, the centrioles move to the opposite poles. The nuclear envelope and the nucleolus disappear and the spindle fibres start appearing. Metaphase is marked by the alignment of chromosomes at the equatorial plate. During anaphase the centromeres divide and the chromatids start moving towards the two opposite poles. Once the chromatids reach the two poles, the chromosomal elongation starts, nucleolus and the nuclear membrane reappear. This stage is called the telophase. Nuclear division is then followed by the cytoplasmic division and is called cytokinesis. Mitosis thus, is the equational division in which the chromosome number of the parent is conserved in the daughter cell. In contrast to mitosis, meiosis occurs in the diploid cells, which are destined to form gametes. It is called the reduction division since it reduces the chromosome number by half while making the gametes. In sexual reproduction when the two gametes fuse the chromosome number is restored to the value in the parent. Meiosis is divided into two phases – meiosis I and meiosis II. In the first meiotic division the homologous chromosomes pair to form bivalents, and undergo crossing over. Meiosis I has a long prophase, which is divided further into five phases. These are leptotene, zygotene, pachytene, diplotene and diakinesis. During metaphase I the bivalents arrange on the equatorial plate. This is followed by anaphase I in which homologous chromosomes move to the opposite poles with both their chromatids. Each pole receives half the chromosome number of the parent cell. In telophase I, the nuclear membrane and nucleolus reappear. Meiosis II is similar to mitosis. During anaphase II the sister chromatids separate. Thus at the end of meiosis four haploid cells are formed. # EXERCISES 1. What is the average cell cycle span for a mammalian cell? 2. Distinguish cytokinesis from karyokinesis. 3. Describe the events taking place during interphase. 4. What is G0 (quiescent phase) of cell cycle?
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# BIOLOGY 1. Why is mitosis called equational division? 2. Name the stage of cell cycle at which one of the following events occur: 1. Chromosomes are moved to spindle equator. 2. Centromere splits and chromatids separate. 3. Pairing between homologous chromosomes takes place. 4. Crossing over between homologous chromosomes takes place. 3. Describe the following: 1. synapsis 2. bivalent 3. chiasmata Draw a diagram to illustrate your answer. 4. How does cytokinesis in plant cells differ from that in animal cells? 5. Find examples where the four daughter cells from meiosis are equal in size and where they are found unequal in size. 6. Distinguish anaphase of mitosis from anaphase I of meiosis. 7. List the main differences between mitosis and meiosis. 8. What is the significance of meiosis? 9. Discuss with your teacher about 1. haploid insects and lower plants where cell-division occurs, and 2. some haploid cells in higher plants where cell-division does not occur. 10. Can there be mitosis without DNA replication in ‘S’ phase? 11. Can there be DNA replication without cell division? 12. Analyse the events during every stage of cell cycle and notice how the following two parameters change 1. number of chromosomes (N) per cell 2. amount of DNA content (C) per cell 2024-25
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# UNIT 2 # STRUCTURAL ORGANISATION IN PLANTS AND ANIMALS # Chapter 5 # Morphology of Flowering Plants # Chapter 6 # Anatomy of Flowering Plants # Chapter 7 # Structural Organisation in Animals The description of the diverse forms of life on earth was made only by observation – through naked eyes or later through magnifying lenses and microscopes. This description is mainly of gross structural features, both external and internal. In addition, observable and perceivable living phenomena were also recorded as part of this description. Before experimental biology or more specifically, physiology, was established as a part of biology, naturalists described only biology. Hence, biology remained as a natural history for a long time. The description, by itself, was amazing in terms of detail. While the initial reaction of a student could be boredom, one should keep in mind that the detailed description was utilised in the later day reductionist biology where living processes drew more attention from scientists than the description of life forms and their structure. Hence, this description became meaningful and helpful in framing research questions in physiology or evolutionary biology. In the following chapters of this unit, the structural organisation of plants and animals, including the structural basis of physiological or behavioural phenomena, is described. For convenience, this description of morphological and anatomical features is presented separately for plants and animals. 2024-25
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Katherine E. Esau was born in Ukraine in 1898. She studied agriculture in Russia and Germany and received her doctorate in 1931 in the United States. She reported in her early publications that the curly top virus spreads through a plant via the food-conducting or phloem tissue. Dr. Esau’s *Plant Anatomy published in 1954 took a dynamic, developmental approach designed to enhance one’s understanding of plant structure and had an enormous impact worldwide, literally bringing about a revival of the discipline. The Anatomy of Seed Plants* by Katherine Esau was published in 1960. It was referred to as Webster’s of plant biology – it is encyclopedic. In 1957 she was elected to the National Academy of Sciences, becoming the sixth woman to receive that honour. In addition to this prestigious award, she received the National Medal of Science from President George Bush in 1989. When Katherine Esau died in the year 1997, Peter Raven, director of Anatomy and Morphology, Missouri Botanical Garden, remembered that she ‘absolutely dominated’ the field of plant biology even at the age of 99. # Katherine Esau (1898 – 1997) # 2024-25
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# CHAPTER 5 # MORPHOLOGY OF FLOWERING PLANTS # 5.1 The Root The wide range in the structure of higher plants will never fail to fascinate us. Even though the angiosperms show such a large diversity in external structure or morphology, they are all characterised by presence of roots, stems, leaves, flowers and fruits. # 5.2 The Stem # 5.3 The Leaf # 5.4 The Inflorescence In chapters 2 and 3, we talked about classification of plants based on morphological and other characteristics. For any successful attempt at classification and at understanding any higher plant (or for that matter any living organism) we need to know standard technical terms and standard definitions. We also need to know about the possible variations in different parts, found as adaptations of the plants to their environment, e.g., adaptations to various habitats, for protection, climbing, storage, etc. # 5.5 The Flower # 5.6 The Fruit # 5.7 The Seed # 5.8 Semi-technical Description of a Typical Flowering Plant If you pull out any weed you will see that all of them have roots, stems and leaves. They may be bearing flowers and fruits. The underground part of the flowering plant is the root system while the portion above the ground forms the shoot system (Figure 5.1). # 5.9 Description of Some Important Families # 5.1 # THE ROOT In majority of the dicotyledonous plants, the direct elongation of the radicle leads to the formation of primary root which grows inside the soil. It bears lateral roots of several orders that are referred to as secondary, tertiary, etc. roots. The primary roots and its branches constitute the
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# BIOLOGY # Flower tap root system, as seen in the mustard plant (Figure 5.2a). In monocotyledonous plants, the primary root is short lived and is replaced by a large number of roots. # Fruit # Stem # Leaf # Shoot system These roots originate from the base of the stem and constitute the fibrous root system, as seen in the wheat plant (Figure 5.2b). In some plants, like grass, Monstera and the banyan tree, roots arise from parts of the plant other than the radicle and are called adventitious roots (Figure 5.2c). The main functions of the root system are absorption of water and minerals from the soil, providing a proper anchorage to the plant parts, storing reserve food material and synthesis of plant growth regulators. # Node # Internode # Bud # Primary root # Root system # Secondary root # Figure 5.1 Parts of a flowering plant Main root Laterals # Figure 5.2 Different types of roots : |(a)|Tap| |---|---| |(b)|Fibrous| |(c)|Adventitious| 2024-25
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dfcd012c-de68-4500-b349-01432aa0aa8d
# MORPHOLOGY OF FLOWERING PLANTS # 5.1.1 Regions of the Root The root is covered at the apex by a thimble-like structure called the root cap (Figure 5.3). It protects the tender apex of the root as it makes its way through the soil. A few millimetres above the root cap is the region of meristematic activity. The cells of this region are very small, thin-walled and with dense protoplasm. They divide repeatedly. The cells proximal to this region undergo rapid elongation and enlargement and are responsible for the growth of the root in length. This region is called the region of elongation. The cells of the elongation zone gradually differentiate and mature. Hence, this zone, proximal to region of elongation, is called the region of maturation. From this region some of the epidermal cells form very fine and delicate, thread-like structures called root hairs. These root hairs absorb water and minerals from the soil. Figure 5.3 The regions of the root-tip # 5.2 THE STEM What are the features that distinguish a stem from a root? The stem is the ascending part of the axis bearing branches, leaves, flowers and fruits. It develops from the plumule of the embryo of a germinating seed. The stem bears nodes and internodes. The region of the stem where leaves are born are called nodes while internodes are the portions between two nodes. The stem bears buds, which may be terminal or axillary. Stem is generally green when young and later often become woody and dark brown. The main function of the stem is spreading out branches bearing leaves, flowers and fruits. It conducts water, minerals and photosynthates. Some stems perform the function of storage of food, support, protection and of vegetative propagation. # 5.3 THE LEAF The leaf is a lateral, generally flattened structure borne on the stem. It develops at the node and bears a bud in its axil. The axillary bud later develops into a branch. Leaves originate from shoot apical meristems and are arranged in an acropetal order. They are the most important vegetative organs for photosynthesis.
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# BIOLOGY # Lamina A typical leaf consists of three main parts: leaf base, petiole and lamina (Figure 5.4 a). The leaf is attached to the stem by the leaf base and may bear two lateral small leaf-like structures called stipules. In monocotyledons, the leaf base expands into a sheath covering the stem partially or wholly. In some leguminous plants, the leaf base may become swollen, which is called the pulvinus. The petiole helps hold the blade to light. Long thin flexible petioles allow leaf blades to flutter in wind, thereby cooling the leaf and bringing fresh air to the leaf surface. The lamina or the leaf blade is the green expanded part of the leaf with veins and veinlets. There is, usually, a middle prominent vein, which is known as the midrib. Veins provide rigidity to the leaf blade and act as channels of transport for water, minerals and food materials. The shape, margin, apex, surface and extent of incision of lamina varies in different leaves. # 5.3.1 Venation The arrangement of veins and the veinlets in the lamina of leaf is termed as venation. When the veinlets form a network, the venation is termed as reticulate (Figure 5.4 b). When the veins run parallel to each other within a lamina, the venation is termed as parallel (Figure 5.4 c). Leaves of dicotyledonous plants generally possess reticulate venation, while parallel venation is the characteristic of most monocotyledons. # 5.3.2 Types of Leaves A leaf is said to be simple when its lamina is entire or when incised, the incisions do not touch the midrib. When the incisions of the lamina reach up to the midrib breaking it into a number of leaflets, the leaf is called compound. A bud is present in the axil of petiole in both simple and compound leaves, but not in the axil of leaflets of the compound leaf. The compound leaves may be of two types (Figure 5.5). In a pinnately compound leaf a (b) palmately compound leaf. # Figure 5.4 Structure of a leaf: - (a) Parts of a leaf - (b) Reticulate venation - (c) Parallel venation # Figure 5.5 Compound leaves: - (a) pinnately compound leaf - (b) palmately compound leaf 2024-25
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# MORPHOLOGY OF FLOWERING PLANTS # 5.3.3 Phyllotaxy Phyllotaxy is the pattern of arrangement of leaves on the stem or branch. This is usually of three types – alternate, opposite and whorled (Figure 5.6). In alternate type of phyllotaxy, a single leaf arises at each node in alternate manner, as in china rose, mustard and sunflower plants. In opposite type, a pair of leaves arise at each node and lie opposite to each other as in Calotropis and guava plants. If more than two leaves arise at a node and form a whorl, it is called whorled, as in Alstonia. # Figure 5.6 Different types of phyllotaxy: - (a) Alternate - (b) Opposite - (c) Whorled # 5.4 THE INFLORESCENCE A flower is a modified shoot wherein the shoot apical meristem changes to floral meristem. Internodes do not elongate and the axis gets condensed. The apex produces different kinds of floral appendages laterally at successive nodes instead of leaves. When a shoot tip transforms into a flower, it is always solitary. The arrangement of flowers on the floral axis is termed as inflorescence. Depending on whether the apex gets developed into a flower or continues to grow, two major types of inflorescences are defined – racemose and cymose. In racemose type of inflorescences the main axis continues to grow, the flowers are borne laterally in an acropetal succession (Figure 5.7). In cymose type of inflorescence the main axis terminates in a flower, hence is limited in growth. The flowers are borne in a basipetal order (Figure 5.7). # Figure 5.7 Racemose inflorescence
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# 5.5 THE FLOWER The flower is the reproductive unit in the angiosperms. It is meant for sexual reproduction. A typical flower has four different kinds of whorls arranged successively on the swollen end of the stalk or pedicel, called thalamus or receptacle. These are calyx, corolla, androecium and gynoecium. Calyx and corolla are accessory organs, while androecium and gynoecium are reproductive organs. In some flowers like lily, the calyx and corolla are not distinct and are termed as perianth. When a flower has both androecium and gynoecium, it is bisexual. A flower having either only stamens or only carpels is unisexual. In symmetry, the flower may be zygomorphic (bilateral symmetry). When a flower can be divided into two equal radial halves in any radial plane passing through the centre, it is said to be actinomorphic, e.g., mustard, datura, chilli. When it can be divided into two similar halves only in one particular vertical plane, it is zygomorphic, e.g., pea, gulmohur, bean, Cassia. A flower is asymmetric (irregular) if it cannot be divided into two similar halves by any vertical plane passing through the centre, as in canna. A flower may be trimerous, tetramerous or pentamerous when the floral appendages are in multiple of 3, 4 or 5, respectively. Flowers with bracts - reduced leaf found at the base of the pedicel - are called bracteate and those without bracts, ebracteate. Figure 5.8 Cymose inflorescence (a) (b) (c) (d) Figure 5.9 Position of floral parts on thalamus: (a) Hypogynous (b) and (c) Perigynous (d) Epigynous
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# MORPHOLOGY OF FLOWERING PLANTS Based on the position of calyx, corolla and androecium in respect of the ovary on thalamus, the flowers are described as hypogynous, perigynous and epigynous (Figure 5.9). In the hypogynous flower the gynoecium occupies the highest position while the other parts are situated below it. The ovary in such flowers is said to be superior, e.g., mustard, china rose and brinjal. If gynoecium is situated in the centre and other parts of the flower are located on the rim of the thalamus almost at the same level, it is called perigynous. The ovary here is said to be half inferior, e.g., plum, rose, peach. In epigynous flowers, the margin of thalamus grows upward enclosing the ovary completely and getting fused with it, the other parts of flower arise above the ovary. Hence, the ovary is said to be inferior as in flowers of guava and cucumber, and the ray florets of sunflower. # 5.5.1 Parts of a Flower Each flower normally has four floral whorls, viz., calyx, corolla, androecium and gynoecium (Figure 5.10). # 5.5.1.1 Calyx The calyx is the outermost whorl of the flower and the members are called sepals. Generally, sepals are green, leaf like and protect the flower in the bud stage. The calyx may be gamosepalous (sepals united) or polysepalous (sepals free). # 5.5.1.2 Corolla Corolla is composed of petals. Petals are usually brightly coloured to attract insects for pollination. Like calyx, corolla may also be gamopetalous (petals united) or polypetalous (petals free). The shape and colour of corolla vary greatly in plants. Corolla may be tubular, bell-shaped, funnel-shaped or wheel-shaped. Aestivation: The mode of arrangement of sepals or petals in floral bud with respect to the other members of the same whorl is known as aestivation. The main types of aestivation are valvate, twisted, imbricate. Androecium Gynoecium Corolla Calyx Pedicel Figure 5.10 Parts of a flower
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# BIOLOGY Figure 5.11 Types of aestivation in corolla: (a) Valvate (b) Twisted (c) Imbricate (d) Vexillary and vexillary (Figure 5.11). When sepals or petals in a whorl just touch one another at the margin, without overlapping, as in Calotropis, it is said to be valvate. If one margin of the appendage overlaps that of the next one and so on as in china rose, lady’s finger and cotton, it is called twisted. If the margins of sepals or petals overlap one another but not in any particular direction as in Cassia and gulmohur, the aestivation is called imbricate. In pea and bean flowers, there are five petals, the largest (standard) overlaps the two lateral petals (wings) which in turn overlap the two smallest anterior petals (keel); this type of aestivation is known as vexillary or papilionaceous. # 5.5.1.3 Androecium Androecium is composed of stamens. Each stamen which represents the male reproductive organ consists of a stalk or a filament and an anther. Each anther is usually bilobed and each lobe has two chambers, the pollen-sacs. The pollen grains are produced in pollen-sacs. A sterile stamen is called staminode. Stamens of flower may be united with other members such as petals or among themselves. When stamens are attached to the petals, they are epipetalous as in brinjal, or epiphyllous when attached to the perianth as in the flowers of lily. The stamens in a flower may either remain free (polyandrous) or may be united in varying degrees. The stamens may be united into one bunch or one bundle or two bundles (diadelphous) as in pea, or into more than two bundles (polyadelphous) as in citrus. There may be a variation in the length of filaments within a flower, as in Salvia and mustard.
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# MORPHOLOGY OF FLOWERING PLANTS # 5.5.1.4 Gynoecium Gynoecium is the female reproductive part of the flower and is made up of one or more carpels. A carpel consists of three parts namely stigma, style and ovary. Ovary is the enlarged basal part, on which lies the elongated tube, the style. The style connects the ovary to the stigma. The stigma is usually at the tip of the style and is the receptive surface for pollen grains. Each ovary bears one or more ovules attached to a flattened, cushion-like placenta. When more than one carpel is present, they may be free (as in lotus and rose) and are called apocarpous. They are termed syncarpous when carpels are fused, as in mustard and tomato. After fertilisation, the ovules develop into seeds and the ovary matures into a fruit. # Placentation The arrangement of ovules within the ovary is known as placentation. The placentation are of different types namely, marginal, axile, parietal, basal, central and free central (Figure 5.12). In marginal placentation the placenta forms a ridge along the ventral suture of the ovary and the ovules are borne on this ridge forming two rows, as in pea. When the placenta is axial and the ovules are attached to it in a multilocular ovary, the placentation is said to be axile, as in china rose, tomato and lemon. In parietal placentation, the ovules develop on the inner wall of the ovary or on peripheral part. Ovary is one-chambered but it becomes two-chambered due to the formation of the false septum, e.g., mustard and Argemone. When the ovules are borne on central axis and septa are absent, as in Dianthus and Primrose the placentation is called free central. In basal placentation, the placenta develops at the base of ovary and a single ovule is attached to it, as in sunflower, marigold. # 5.6 THE FRUIT The fruit is a characteristic feature of the flowering plants. It is a mature or ripened ovary, developed after fertilisation. If a fruit is formed without fertilisation of the ovary, it is called a parthenocarpic fruit. Generally, the fruit consists of a wall or pericarp and seeds. The pericarp may be dry or fleshy. When pericarp is thick and fleshy, it is differentiated into the outer epicarp, the middle mesocarp and the inner endocarp. In mango and coconut, the fruit is known as a drupe (Figure 5.13). They develop from monocarpellary superior ovaries and are one seeded. In mango the pericarp is well differentiated into an. # Figure 5.12 Types of placentation: (a) Marginal (b) Axile (c) Parietal (d) Free central (e) Basal
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# 5.7 THE SEED The ovules after fertilisation, develop into seeds. A seed is made up of a seed coat and an embryo. The embryo is made up of a radicle, an embryonal axis and one (as in wheat, maize) or two cotyledons (as in gram and pea). # 5.7.1 Structure of a Dicotyledonous Seed The outermost covering of a seed is the seed coat. The seed coat has two layers, the outer testa and the inner tegmen. The hilum is a scar on the seed coat through which the developing seeds were attached to the fruit. Above the hilum is a small pore called the micropyle. Within the seed coat is the embryo, consisting of an embryonal axis and two cotyledons. The cotyledons are often fleshy and full of reserve food materials. At the two ends of the embryonal axis are present the radicle and the plumule (Figure 5.14). In some seeds such as castor the endosperm formed as a result of double fertilisation, is a food storing tissue and called endospermic seeds. In plants such as bean, gram and pea, the endosperm is not present in mature seeds and such seeds are called non-endospermous. # 5.7.2 Structure of Monocotyledonous Seed Generally, monocotyledonous seeds are endospermic but some as in orchids are non-endospermic. In the seeds of cereals such as maize the |Figure 5.13 Parts of a fruit|Figure 5.13 Parts of a fruit| |---| |(a) Mango|(b) Coconut| |Figure 5.14 Structure of dicotyledonous seed|Figure 5.14 Structure of dicotyledonous seed| |---| |Seed coat|Hilum| |Cotyledon|Radicle| |Plumule|Micropyle|
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# MORPHOLOGY OF FLOWERING PLANTS # 5.8 SEMI-TECHNICAL DESCRIPTION OF A TYPICAL FLOWERING PLANT Various morphological features are used to describe a flowering plant. The description has to be brief, in a simple and scientific language and presented in a proper sequence. The plant is described beginning with its habit, vegetative characters – roots, stem and leaves and then floral characters inflorescence and flower parts. After describing various parts of plant, a floral diagram and a floral formula are presented. The floral formula is represented by some symbols. In the floral formula, Br stands for bracteate, K stands for calyx, C for corolla, P for perianth, A for androecium and G for Gynoecium, G ⊕ K2+2 C A2+4 G(2) 4 for superior ovary and G for inferior ovary, for male, for female, for bisexual plants, ⊕ for actinomorphic. # Figure 5.15 Structure of a monocotyledonous seed |Seed coat & fruit-wall|Endosperm| |---|---| |Aleurone layer|Scutellum| | |Coleoptile| |Endosperm|Plumule| |Embryo|Radicle| | |Coleorhiza| The seed coat is membranous and generally fused with the fruit wall. The endosperm is bulky and stores food. The outer covering of endosperm separates the embryo by a proteinous layer called aleurone layer. The embryo is small and situated in a groove at one end of the endosperm. It consists of one large and shield shaped cotyledon known as scutellum and a short axis with a plumule and a radicle. The plumule and radicle are enclosed in sheaths which are called coleoptile and coleorhiza respectively (Figure 5.15). # Figure 5.16 Floral diagram with floral formula
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# 68 # BIOLOGY and for zygomorphic nature of flower. Fusion is indicated by enclosing the figure within bracket and adhesion by a line drawn above the symbols of the floral parts. A floral diagram provides information about the number of parts of a flower, their arrangement and the relation they have with one another (Figure 5.16). The position of the mother axis with respect to the flower is represented by a dot on the top of the floral diagram. Calyx, corolla, androecium and gynoecium are drawn in successive whorls, calyx being the outermost and the gynoecium being in the centre. Floral formula also shows cohesion and adhesion within parts of whorls and between whorls. The floral diagram and floral formula in Figure 5.16 represents the mustard plant (Family: Brassicaceae). # 5.9 # SOLANACEAE It is a large family, commonly called as the ‘potato family’. It is widely distributed in tropics, subtropics and even temperate zones (Figure 5.17). # Vegetative Characters Plants mostly herbs, shrubs and rarely small trees Stem: herbaceous rarely woody, aerial; erect, cylindrical, branched, solid or hollow, hairy or glabrous, underground stem in potato (Solanum tuberosum) Leaves: alternate, simple, rarely pinnately compound, exstipulate; venation reticulate # Figure 5.17 Solanum nigrum (makoi) plant: (a) Flowering twig (b) Flower (c) L.S. of flower (d) Stamens (e) Carpel (f) Floral diagram 2024-25
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# MORPHOLOGY OF FLOWERING PLANTS # Floral Characters - Inflorescence: Solitary, axillary or cymose as in Solanum - Flower: bisexual, actinomorphic - Calyx: sepals five, united, persistent, valvate aestivation - Corolla: petals five, united; valvate aestivation - Androecium: stamens five, epipetalous - Gynoecium: bicarpellary obligately placed, syncarpous; ovary superior bilocular, placenta swollen with many ovules, axile - Fruits: berry or capsule - Seeds: many, endospermous - Floral Formula: ⊕ # Economic Importance Many plants belonging to this family are source of food (tomato, brinjal, potato), spice (chilli); medicine (belladonna, ashwagandha, tobacco); ornamentals (petunia). # SUMMARY Flowering plants exhibit enormous variation in shape, size, structure, mode of nutrition, life span, habit and habitat. They have well developed root and shoot systems. Root system is either tap root or fibrous. Generally, dicotyledonous plants have tap roots while monocotyledonous plants have fibrous roots. The roots in some plants get modified for storage of food, mechanical support and respiration. The shoot system is differentiated into stem, leaves, flowers and fruits. The morphological features of stems like the presence of nodes and internodes, multicellular hair and positively phototropic nature help to differentiate the stems from roots. Leaf is a lateral outgrowth of stem developed exogeneously at the node. These are green in colour to perform the function of photosynthesis. Leaves exhibit marked variations in their shape, size, margin, apex and extent of incisions of leaf blade (lamina). The flower is a modified shoot, meant for sexual reproduction. The flowers are arranged in different types of inflorescences. They exhibit enormous variation in structure, symmetry, position of ovary in relation to other parts, arrangement of petals, sepals, ovules etc. After fertilisation, the ovary is modified into fruits and ovules into seeds. Seeds either may be monocotyledonous or dicotyledonous. They vary in shape, size and period of viability. The floral characteristics form the basis of classification.
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# BIOLOGY and identification of flowering plants. This can be illustrated through semi-technical descriptions of families. Hence, a flowering plant is described in a definite sequence by using scientific terms. The floral features are represented in the summarised form as floral diagrams and floral formula. # EXERCISES 1. How is a pinnately compound leaf different from a palmately compound leaf? 2. Explain with suitable examples the different types of phyllotaxy. 3. Define the following terms: - (a) aestivation - (b) placentation - (c) actinomorphic - (d) zygomorphic - (e) superior ovary - (f) perigynous flower - (g) epipetalous stamen 4. Differentiate between - (a) Racemose and cymose inflorescence - (b) Apocarpous and syncarpous ovary 5. Draw the labelled diagram of the following: - (i) gram seed - (ii) V.S. of maize seed 6. Take one flower of the family Solanaceae and write its semi-technical description. Also draw their floral diagram. 7. Describe the various types of placentations found in flowering plants. 8. What is a flower? Describe the parts of a typical angiosperm flower. 9. Define the term inflorescence. Explain the basis for the different types of inflorescence in flowering plants. 10. Describe the arrangement of floral members in relation to their insertion on thalamus. 2024-25
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# CHAPTER 19 # C HEMICAL C OORDINATION AND I NTEGRATION # 19.1 Endocrine Glands and Hormones You have already learnt that the neural system provides a point-to-point rapid coordination among organs. The neural coordination is fast but short-lived. As the nerve fibres do not innervate all cells of the body and the cellular functions need to be continuously regulated; a special kind of coordination and integration has to be provided. This function is carried out by hormones. The neural system and the endocrine system jointly coordinate and regulate the physiological functions in the body. # 19.2 Human Endocrine System # 19.3 Hormones of Heart, Kidney and Gastrointestinal Tract # 19.4 Mechanism of Hormone Action Endocrine glands lack ducts and are hence, called ductless glands. Their secretions are called hormones. The classical definition of hormone as a chemical produced by endocrine glands and released into the blood and transported to a distantly located target organ has current scientific definition as follows: Hormones are non-nutrient chemicals which act as intercellular messengers and are produced in trace amounts. The new definition covers a number of new molecules in addition to the hormones secreted by the organised endocrine glands. Invertebrates possess very simple endocrine systems with few hormones whereas a large number of chemicals act as hormones and provide coordination in the vertebrates. The human endocrine system is described here.
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# 19.2 HUMAN ENDOCRINE SYSTEM The endocrine glands and hormone producing diffused tissues/cells located in different parts of our body constitute the endocrine system. Pituitary, pineal, thyroid, adrenal, pancreas, parathyroid, thymus and gonads (testis in males and ovary in females) are the organised endocrine bodies in our body (Figure 19.1). In addition to these, some other organs, e.g., gastrointestinal tract, liver, kidney, heart also produce hormones. A brief account of the structure and functions of all major endocrine glands and hypothalamus of the human body is given in the following sections. # 19.2.1 The Hypothalamus As you know, the hypothalamus is the basal part of diencephalon, forebrain (Figure 19.1) and it regulates a wide spectrum of body functions. It contains several groups of neurosecretory cells called nuclei which produce hormones. These hormones regulate the synthesis and secretion of pituitary hormones. However, the hormones produced by hypothalamus are of two types, the releasing hormones (which stimulate secretion of pituitary hormones) and the inhibiting hormones (which inhibit secretions of pituitary hormones). For example a hypothalamic hormone called Gonadotrophin releasing hormone (GnRH) stimulates the pituitary synthesis and release of gonadotrophins. On the other hand, somatostatin from the hypothalamus inhibits the release of growth hormone from the pituitary. These hormones originating in the hypothalamic neurons, pass through axons and are released from their nerve endings. These hormones reach the pituitary gland through a portal circulatory system and regulate the functions of the anterior pituitary. The posterior pituitary is under the direct neural regulation of the hypothalamus (Figure 19.2).
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# 19.2.2 The Pituitary Gland The pituitary gland is located in a bony cavity called sella tursica and is attached to hypothalamus by a stalk (Figure 19.2). It is divided anatomically into an adenohypophysis and a neurohypophysis. Adenohypophysis consists of two portions, pars distalis and pars intermedia. The pars distalis region of pituitary, commonly called anterior pituitary, produces growth hormone (GH), prolactin (PRL), thyroid stimulating hormone (TSH), adrenocorticotrophic hormone (ACTH), luteinizing hormone (LH) and follicle stimulating hormone (FSH). Pars intermedia secretes only one hormone called melanocyte stimulating hormone (MSH). However, in humans, the pars intermedia is almost merged with pars distalis. Neurohypophysis (pars nervosa) also known as posterior pituitary, stores and releases two hormones called oxytocin and vasopressin, which are actually synthesised by the hypothalamus and are transported axonally to neurohypophysis. Over-secretion of GH stimulates abnormal growth of the body leading to gigantism and low secretion of GH results in stunted growth resulting in pituitary dwarfism. Excess secretion of growth hormone in adults especially in middle age can result in severe disfigurement (especially of the face) called Acromegaly, which may lead to serious complications, and premature death if unchecked. The disease is hard to diagnose in the early stages and often goes undetected for many years, until changes in external features become noticeable. Prolactin regulates the growth of the mammary glands and formation of milk in them. TSH stimulates the synthesis and secretion of thyroid hormones from the thyroid gland. ACTH stimulates the synthesis and secretion of steroid hormones called glucocorticoids from the adrenal cortex. LH and FSH stimulate gonadal activity and hence are called gonadotrophins. In males, LH stimulates the synthesis and secretion of hormones called androgens from testis. In males, FSH and androgens regulate spermatogenesis. In females, LH induces ovulation of fully mature follicles (graafian follicles) and maintains the corpus luteum, formed from the remnants of the graafian follicles after ovulation. FSH stimulates growth and development of the ovarian.
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# 19.2.3 The Pineal Gland The pineal gland is located on the dorsal side of forebrain. Pineal secretes a hormone called melatonin. Melatonin plays a very important role in the regulation of a 24-hour (diurnal) rhythm of our body. For example, it helps in maintaining the normal rhythms of sleep-wake cycle, body temperature. In addition, melatonin also influences metabolism, pigmentation, the menstrual cycle as well as our defense capability. # 19.2.4 Thyroid Gland The thyroid gland is composed of two lobes which are located on either side of the trachea (Figure 19.3 a). Both the lobes are interconnected with a thin flap of connective tissue called isthmus. The thyroid gland is composed of follicles and stromal tissues. Each thyroid follicle is composed of follicular cells, enclosing a cavity. These follicular cells synthesise two hormones, tetraiodothyronine or thyroxine (T4) and triiodothyronine (T3). Iodine is essential for the normal rate of hormone synthesis in the thyroid. Deficiency of iodine in our diet results in hypothyroidism and goitre. Hypothyroidism during pregnancy causes defective development and maturation of the growing baby leading to stunted growth (cretinism), mental. # Figure 19.3 Diagrammatic view of the enlargement of the thyroid gland, commonly called position of Thyroid and Parathyroid (a) Ventral side (b) Dorsal side
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# Chemical Coordination and Integration retardation, low intelligence quotient, abnormal skin, deaf-mutism, etc. In adult women, hypothyroidism may cause menstrual cycle to become irregular. Due to cancer of the thyroid gland or due to development of nodules of the thyroid glands, the rate of synthesis and secretion of the thyroid hormones is increased to abnormal high levels leading to a condition called hyperthyroidism which adversely affects the body physiology. Exopthalmic goitre is a form of hyperthyroidism, characterised by enlargement of the thyroid gland, protrusion of the eyeballs, increased basal metabolic rate, and weight loss, also called Graves’ disease. Thyroid hormones play an important role in the regulation of the basal metabolic rate. These hormones also support the process of red blood cell formation. Thyroid hormones control the metabolism of carbohydrates, proteins and fats. Maintenance of water and electrolyte balance is also influenced by thyroid hormones. Thyroid gland also secretes a protein hormone called thyrocalcitonin (TCT) which regulates the blood calcium levels. # 19.2.5 Parathyroid Gland In humans, four parathyroid glands are present on the back side of the thyroid gland, one pair each in the two lobes of the thyroid gland (Figure 19.3 b). The parathyroid glands secrete a peptide hormone called parathyroid hormone (PTH). The secretion of PTH is regulated by the circulating levels of calcium ions. Parathyroid hormone (PTH) increases the Ca2+ levels in the blood. PTH acts on bones and stimulates the process of bone resorption (dissolution/demineralisation). PTH also stimulates reabsorption of Ca2+ by the renal tubules and increases Ca2+ absorption from the digested food. It is, thus, clear that PTH is a hypercalcemic hormone, i.e., it increases the blood Ca2+ levels. Along with TCT, it plays a significant role in calcium balance in the body. # 19.2.6 Thymus The thymus gland is a lobular structure located between lungs behind sternum on the ventral side of aorta. The thymus plays a major role in the development of the immune system. This gland secretes the peptide hormones called thymosins. Thymosins play a major role in the differentiation of T-lymphocytes, which provide cell-mediated immunity. In addition, thymosins also promote production of antibodies to provide humoral immunity. Thymus is degenerated in old individuals resulting in a decreased production of thymosins. As a result, the immune responses of old persons become weak.
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# 19.2.7 Adrenal Gland Our body has one pair of adrenal glands, one at the anterior part of each kidney (Figure 19.4 a). The gland is composed of two types of tissues. The centrally located tissue is called the adrenal medulla, and outside this lies the adrenal cortex (Figure 19.4 b). Underproduction of hormones by the adrenal cortex alters carbohydrate metabolism causing acute weakness and fatigue leading to a disease called Addison’s disease. |Adrenal gland|Adrenal cortex| |---|---| |Adrenal medulla|Kidney| |(a)| | Figure 19.4 Diagrammatic representation of: (a) Adrenal gland above kidney (b) Section showing two parts of adrenal gland The adrenal medulla secretes two hormones called adrenaline or epinephrine and noradrenaline or norepinephrine. These are commonly called catecholamines. Adrenaline and noradrenaline are rapidly secreted in response to stress of any kind and during emergency situations and are called emergency hormones or hormones of Fight or Flight. These hormones increase alertness, pupilary dilation, piloerection (raising of hairs), sweating etc. Both the hormones increase the heart beat, the strength of heart contraction and the rate of respiration. Catecholamines also stimulate the breakdown of glycogen resulting in
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# Chemical Coordination and Integration an increased concentration of glucose in blood. In addition, they also stimulate the breakdown of lipids and proteins. The adrenal cortex can be divided into three layers, called zona reticularis (inner layer), zona fasciculata (middle layer) and zona glomerulosa (outer layer). The adrenal cortex secretes many hormones, commonly called as corticoids. The corticoids, which are involved in carbohydrate metabolism are called glucocorticoids. In our body, cortisol is the main glucocorticoid. Corticoids, which regulate the balance of water and electrolytes in our body are called mineralocorticoids. Aldosterone is the main mineralocorticoid in our body. Glucocorticoids stimulate gluconeogenesis, lipolysis and proteolysis; and inhibit cellular uptake and utilisation of amino acids. Cortisol is also involved in maintaining the cardio-vascular system as well as the kidney functions. Glucocorticoids, particularly cortisol, produces anti-inflammatory reactions and suppresses the immune response. Cortisol stimulates the RBC production. Aldosterone acts mainly at the renal tubules and stimulates the reabsorption of Na and water and excretion of K+ and phosphate ions. Thus, aldosterone helps in the maintenance of electrolytes, body fluid volume, osmotic pressure and blood pressure. Small amounts of androgenic steroids are also secreted by the adrenal cortex which play a role in the growth of axial hair, pubic hair and facial hair during puberty. # 19.2.8 Pancreas Pancreas is a composite gland (Figure 19.1) which acts as both exocrine and endocrine gland. The endocrine pancreas consists of ‘Islets of Langerhans’. There are about 1 to 2 million Islets of Langerhans in a normal human pancreas representing only 1 to 2 per cent of the pancreatic tissue. The two main types of cells in the Islet of Langerhans are called α-cells and β-cells. The α-cells secrete a hormone called glucagon, while the β-cells secrete insulin. Glucagon is a peptide hormone, and plays an important role in maintaining the normal blood glucose levels. Glucagon acts mainly on the liver cells (hepatocytes) and stimulates glycogenolysis resulting in an increased blood sugar (hyperglycemia). In addition, this hormone stimulates the process of gluconeogenesis which also contributes to hyperglycemia. Glucagon reduces the cellular glucose uptake and utilisation. Thus, glucagon is a hyperglycemic hormone. Insulin is a peptide hormone, which plays a major role in the regulation of glucose homeostasis. Insulin acts mainly on hepatocytes and adipocytes (cells of adipose tissue), and enhances cellular glucose.
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# BIOLOGY uptake and utilisation. As a result, there is a rapid movement of glucose from blood to hepatocytes and adipocytes resulting in decreased blood glucose levels (hypoglycemia). Insulin also stimulates conversion of glucose to glycogen (glycogenesis) in the target cells. The glucose homeostasis in blood is thus maintained jointly by the two – insulin and glucagons. Prolonged hyperglycemia leads to a complex disorder called diabetes mellitus which is associated with loss of glucose through urine and formation of harmful compounds known as ketone bodies. Diabetic patients are successfully treated with insulin therapy. # 19.2.9 Testis A pair of testis is present in the scrotal sac (outside abdomen) of male individuals (Figure 19.1). Testis performs dual functions as a primary sex organ as well as an endocrine gland. Testis is composed of seminiferous tubules and stromal or interstitial tissue. The Leydig cells or interstitial cells, which are present in the intertubular spaces produce a group of hormones called androgens mainly testosterone. Androgens regulate the development, maturation and functions of the male accessory sex organs like epididymis, vas deferens, seminal vesicles, prostate gland, urethra etc. These hormones stimulate muscular growth, growth of facial and axillary hair, aggressiveness, low pitch of voice etc. Androgens play a major stimulatory role in the process of spermatogenesis (formation of spermatozoa). Androgens act on the central neural system and influence the male sexual behaviour (libido). These hormones produce anabolic (synthetic) effects on protein and carbohydrate metabolism. # 19.2.10 Ovary Females have a pair of ovaries located in the abdomen (Figure 19.1). Ovary is the primary female sex organ which produces one ovum during each menstrual cycle. In addition, ovary also produces two groups of steroid hormones called estrogen and progesterone. Ovary is composed of ovarian follicles and stromal tissues. The estrogen is synthesised and secreted mainly by the growing ovarian follicles. After ovulation, the ruptured follicle is converted to a structure called corpus luteum, which secretes mainly progesterone. Estrogens produce wide ranging actions such as stimulation of growth and activities of female secondary sex organs, development of growing
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# Chemical Coordination and Integration Ovarian follicles, appearance of female secondary sex characters (e.g., high pitch of voice, etc.), mammary gland development. Estrogens also regulate female sexual behaviour. Progesterone supports pregnancy. Progesterone also acts on the mammary glands and stimulates the formation of alveoli (sac-like structures which store milk) and milk secretion. # 19.3 Hormones of Heart, Kidney and Gastrointestinal Tract Now you know about the endocrine glands and their hormones. However, as mentioned earlier, hormones are also secreted by some tissues which are not endocrine glands. For example, the atrial wall of our heart secretes a very important peptide hormone called atrial natriuretic factor (ANF), which decreases blood pressure. When blood pressure is increased, ANF is secreted which causes dilation of the blood vessels. This reduces the blood pressure. The juxtaglomerular cells of kidney produce a peptide hormone called erythropoietin which stimulates erythropoiesis (formation of RBC). Endocrine cells present in different parts of the gastro-intestinal tract secrete four major peptide hormones, namely gastrin, secretin, cholecystokinin (CCK) and gastric inhibitory peptide (GIP). Gastrin acts on the gastric glands and stimulates the secretion of hydrochloric acid and pepsinogen. Secretin acts on the exocrine pancreas and stimulates secretion of water and bicarbonate ions. CCK acts on both pancreas and gall bladder and stimulates the secretion of pancreatic enzymes and bile juice, respectively. GIP inhibits gastric secretion and motility. Several other non-endocrine tissues secrete hormones called growth factors. These factors are essential for the normal growth of tissues and their repairing/regeneration. # 19.4 Mechanism of Hormone Action Hormones produce their effects on target tissues by binding to specific proteins called hormone receptors located in the target tissues only. Hormone receptors present on the cell membrane of the target cells are called membrane-bound receptors and the receptors present inside the target cell are called intracellular receptors, mostly nuclear receptors (present in the nucleus). Binding of a hormone to its receptor leads to the formation of a hormone-receptor complex. Each receptor is specific to one hormone only and hence receptors are specific. Hormone-Receptor complex formation leads to certain biochemical changes in the target tissue. Target tissue metabolism and hence
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# BIOLOGY Physiological functions are regulated by hormones. On the basis of their chemical nature, hormones can be divided into groups: 1. Peptide, polypeptide, protein hormones (e.g., insulin, glucagon, pituitary hormones, hypothalamic hormones, etc.) 2. Steroids (e.g., cortisol, testosterone, estradiol and progesterone) 3. Iodothyronines (thyroid hormones) 4. Amino-acid derivatives (e.g., epinephrine). Hormones which interact with membrane-bound receptors normally do not enter the target cell, but generate second messengers (e.g., cyclic AMP, IP, Ca++ etc.) which in turn regulate cellular metabolism (Figure 19.5a). Hormones which interact with intracellular receptors (e.g., steroid hormones, iodothyronines, etc.) mostly regulate gene expression or chromosome function by the interaction of hormone-receptor complex with the genome. Cumulative biochemical actions result in physiological and developmental effects (Figure 19.5b). (a) 2024-25
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# Chemical Coordination and Integration # Figure 19.5 Diagramatic representation of the mechanism of hormone action: - (a) Protein hormone - (b) Steroid hormone # Summary There are special chemicals which act as hormones and provide chemical coordination, integration and regulation in the human body. These hormones regulate metabolism, growth and development of our organs, the endocrine glands or certain cells. The endocrine system is composed of hypothalamus, pituitary and pineal, thyroid, adrenal, pancreas, parathyroid, thymus and gonads (testis and ovary). In addition to these, some other organs, e.g., gastrointestinal tract, kidney, heart etc., also produce hormones. The pituitary gland is divided into three major parts, which are called as pars distalis, pars intermedia and pars nervosa. Pars distalis produces six trophic hormones. Pars intermedia secretes
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# BIOLOGY only one hormone, while pars nervosa (neurohypophysis) secretes two hormones. The pituitary hormones regulate the growth and development of somatic tissues and activities of peripheral endocrine glands. Pineal gland secretes melatonin, which plays a very important role in the regulation of 24-hour (diurnal) rhythms of our body (e.g., rhythms of sleep and state of being awake, body temperature, etc.). The thyroid gland hormones play an important role in the regulation of the basal metabolic rate, development and maturation of the central neural system, erythropoiesis, metabolism of carbohydrates, proteins and fats, menstrual cycle. Another thyroid hormone, i.e., thyrocalcitonin regulates calcium levels in our blood by decreasing it. The parathyroid glands secrete parathyroid hormone (PTH) which increases the blood Ca2+ levels and plays a major role in calcium homeostasis. The thymus gland secretes thymosins which play a major role in the differentiation of T-lymphocytes, which provide cell-mediated immunity. In addition, thymosins also increase the production of antibodies to provide humoral immunity. The adrenal gland is composed of the centrally located adrenal medulla and the outer adrenal cortex. The adrenal medulla secretes epinephrine and norepinephrine. These hormones increase alertness, pupilary dilation, piloerection, sweating, heart beat, strength of heart contraction, rate of respiration, glycogenolysis, lipolysis, proteolysis. The adrenal cortex secretes glucocorticoids and mineralocorticoids. Glucocorticoids stimulate gluconeogenesis, lipolysis, proteolysis, erythropoiesis, cardio-vascular system, blood pressure, and glomerular filtration rate and inhibit inflammatory reactions by suppressing the immune response. Mineralocorticoids regulate water and electrolyte contents of the body. The endocrine pancreas secretes glucagon and insulin. Glucagon stimulates glycogenolysis and gluconeogenesis resulting in hyperglycemia. Insulin stimulates cellular glucose uptake and utilisation, and glycogenesis resulting in hypoglycemia. Insulin deficiency and/or insulin resistance result in a disease called diabetes mellitus. The testis secretes androgens, which stimulate the development, maturation and functions of the male accessory sex organs, appearance of the male secondary sex characters, spermatogenesis, male sexual behaviour, anabolic pathways and erythropoiesis. The ovary secretes estrogen and progesterone. Estrogen stimulates growth and development of female accessory sex organs and secondary sex characters. Progesterone plays a major role in the maintenance of pregnancy as well as in mammary gland development and lactation. The atrial wall of the heart produces atrial natriuretic factor which decreases the blood pressure. Kidney produces erythropoietin which stimulates erythropoiesis. The gastrointestinal tract secretes gastrin, secretin, cholecystokinin and gastric inhibitory peptide. These hormones regulate the secretion of digestive juices and help in digestion. 2024-25
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# Chemical Coordination and Integration # Exercises 1. Define the following: 1. Exocrine gland 2. Endocrine gland 3. Hormone 2. Diagrammatically indicate the location of the various endocrine glands in our body. 3. List the hormones secreted by the following: 1. Hypothalamus 2. Pituitary 3. Thyroid 4. Parathyroid 5. Adrenal 6. Pancreas 7. Testis 8. Ovary 9. Thymus 10. Atrium 11. Kidney 12. G-I Tract 4. Fill in the blanks: |Hormones|Target gland| |---|---| |Hypothalamic hormones|__________________| |Thyrotrophin (TSH)|__________________| |Corticotrophin (ACTH)|__________________| |Gonadotrophins (LH, FSH)|__________________| |Melanotrophin (MSH)|__________________| 5. Write short notes on the functions of the following hormones: 1. Parathyroid hormone (PTH) 2. Thyroid hormones 3. Thymosins 4. Androgens 5. Estrogens 6. Insulin and Glucagon 6. Give example(s) of: 1. Hyperglycemic hormone and hypoglycemic hormone 2. Hypercalcemic hormone 3. Gonadotrophic hormones 4. Progestational hormone 5. Blood pressure lowering hormone 6. Androgens and estrogens 7. Which hormonal deficiency is responsible for the following: 1. Diabetes mellitus 2. Goitre 3. Cretinism 8. Briefly mention the mechanism of action of FSH. 9. Match the following: |Column I|Column II| |---|---| |(a) T4|(i) Hypothalamus| |(b) PTH|(iv) Parathyroid| |(c) GnRH|(iii) Pituitary| |(d) LH|(ii) Thyroid|
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NOTE 2024-25
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# CHAPTER 18 # NEURAL CONTROL AND COORDINATION # 18.1 Neural System As you know, the functions of the organs/organ systems in our body must be coordinated to maintain homeostasis. Coordination is the process through which two or more organs interact and complement the functions of one another. For example, when we do physical exercises, the energy demand is increased for maintaining an increased muscular activity. The supply of oxygen is also increased. The increased supply of oxygen necessitates an increase in the rate of respiration, heart beat and increased blood flow via blood vessels. When physical exercise is stopped, the activities of nerves, lungs, heart and kidney gradually return to their normal conditions. Thus, the functions of muscles, lungs, heart, blood vessels, kidney and other organs are coordinated while performing physical exercises. In our body the neural system and the endocrine system jointly coordinate and integrate all the activities of the organs so that they function in a synchronised fashion. # 18.2 Human Neural System The neural system provides an organised network of point-to-point connections for a quick coordination. The endocrine system provides chemical integration through hormones. In this chapter, you will learn about the neural system of human, mechanisms of neural coordination like transmission of nerve impulse, impulse conduction across a synapse.
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# N EURAL CONTROL AND COORDINATION # 18.1 N EURAL SYSTEM The neural system of all animals is composed of highly specialised cells called neurons which can detect, receive and transmit different kinds of stimuli. The neural organisation is very simple in lower invertebrates. For example, in Hydra it is composed of a network of neurons. The neural system is better organised in insects, where a brain is present along with a number of ganglia and neural tissues. The vertebrates have a more developed neural system. # 18.2 H UMAN N EURAL SYSTEM The human neural system is divided into two parts: 1. the central neural system (CNS) 2. the peripheral neural system (PNS) The CNS includes the brain and the spinal cord and is the site of information processing and control. The PNS comprises of all the nerves of the body associated with the CNS (brain and spinal cord). The nerve fibres of the PNS are of two types: - (a) afferent fibres - (b) efferent fibres The afferent nerve fibres transmit impulses from tissues/organs to the CNS and the efferent fibres transmit regulatory impulses from the CNS to the concerned peripheral tissues/organs. The PNS is divided into two divisions called somatic neural system and autonomic neural system. The somatic neural system relays impulses from the CNS to skeletal muscles while the autonomic neural system transmits impulses from the CNS to the involuntary organs and smooth muscles of the body. The autonomic neural system is further classified into sympathetic neural system and parasympathetic neural system. Visceral nervous system is the part of the peripheral nervous system that comprises the whole complex of nerves, fibres, ganglia, and plexuses by which impulses travel from the central nervous system to the viscera and from the viscera to the central nervous system. # 18.3 N EURON AS STRUCTURAL AND FUNCTIONAL UNIT OF N EURAL SYSTEM A neuron is a microscopic structure composed of three major parts, namely, cell body, dendrites and axon (Figure 18.1). The cell body contains cytoplasm with typical cell organelles and certain granular bodies called Nissl’s granules. Short fibres which branch repeatedly and project out of the cell body also.
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# BIOLOGY contain Nissl’s granules and are called dendrites. These fibres transmit impulses towards the cell body. The axon is a long fibre, the distal end of which is branched. Each branch terminates as a bulb-like structure called synaptic knob which possess synaptic vesicles containing chemicals called neurotransmitters. The axons transmit nerve impulses away from the cell body to a synapse or to a neuro-muscular junction. Based on the number of axon and dendrites, the neurons are divided into three types, i.e., multipolar (with one axon and two or more dendrites; found in the cerebral cortex), bipolar (with one axon and one dendrite, found in the retina of eye) and unipolar (cell body with one axon only; found usually in the embryonic stage). There are two types of axons, namely, myelinated and non-myelinated. The myelinated nerve fibres are enveloped with Schwann cells, which form a myelin sheath around the axon. The gaps between two adjacent myelin sheaths are called nodes of Ranvier. Myelinated nerve fibres are found in spinal and cranial nerves. Unmyelinated nerve fibre is enclosed by a Schwann cell that does not form a myelin sheath around the axon, and is commonly found in autonomous and the somatic neural systems. # 18.3.1 Generation and Conduction of Nerve Impulse Neurons are excitable cells because their membranes are in a polarised state. Do you know why the membrane of a neuron is polarised? Different types of ion channels are present on the neural membrane. These ion channels are selectively permeable to different ions. When a neuron is not conducting any impulse, i.e., resting, the axonal membrane is comparatively more permeable to potassium ions (K+) and nearly impermeable to sodium ions (Na+). Similarly, the membrane is impermeable to negatively charged proteins present in the axoplasm. Consequently, the axoplasm inside the axon contains high concentration of K+ and negatively charged proteins and low concentration of Na+. In contrast, the fluid outside the axon contains a low concentration of K+, a high concentration of Na+ and thus form a concentration gradient. These ionic gradients across the resting membrane are maintained by the active transport of ions by the sodium-potassium pump which transports 3 Na+ outwards for 2 K+ into the cell. As a result, the outer surface of the axonal membrane possesses a positive charge while its inner surface
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# N EURAL CONTROL AND COORDINATION # Figure 18.2 Diagrammatic representation of impulse conduction through an axon (at points A and B) becomes negatively charged and therefore is polarised. The electrical potential difference across the resting plasma membrane is called as the resting potential. You might be curious to know about the mechanisms of generation of nerve impulse and its conduction along an axon. When a stimulus is applied at a site (Figure 18.2 e.g., point A) on the polarised membrane, the membrane at the site A becomes freely permeable to Na+. This leads to a rapid influx of Na+ followed by the reversal of the polarity at that site, i.e., the outer surface of the membrane becomes negatively charged and the inner side becomes positively charged. The polarity of the membrane at the site A is thus reversed and hence depolarised. The electrical potential difference across the plasma membrane at the site A is called the action potential, which is in fact termed as a nerve impulse. At sites immediately ahead, the axon (e.g., site B) membrane has a positive charge on the outer surface and a negative charge on its inner surface. As a result, a current flows on the inner surface from site A to site B. On the outer surface current flows from site B to site A (Figure 18.2) to complete the circuit of current flow. Hence, the polarity at the site is reversed, and an action potential is generated at site B. Thus, the impulse (action potential) generated at site A arrives at site B. The sequence is repeated along the length of the axon and consequently the impulse is conducted. The rise in the stimulus-induced permeability to Na+ is extremely short-lived. It is quickly followed by a rise in permeability to K. Within a fraction of a second, K+ diffuses outside the membrane and restores the resting potential of the membrane at the site of excitation and the fibre becomes once more responsive to further stimulation.
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# 18.3.2 Transmission of Impulses A nerve impulse is transmitted from one neuron to another through junctions called synapses. A synapse is formed by the membranes of a pre-synaptic neuron and a post-synaptic neuron, which may or may not be separated by a gap called synaptic cleft. There are two types of synapses, namely, electrical synapses and chemical synapses. At electrical synapses, the membranes of pre- and post-synaptic neurons are in very close proximity. Electrical current can flow directly from one neuron into the other across these synapses. Transmission of an impulse across electrical synapses is very similar to impulse conduction along a single axon. Impulse transmission across an electrical synapse is always faster than that across a chemical synapse. Electrical synapses are rare in our system. At a chemical synapse, the membranes of the pre- and post-synaptic neurons are separated by a fluid-filled space called synaptic cleft (Figure 18.3). Do you know how the pre-synaptic neuron transmits an impulse (action potential) across the synaptic cleft to the post-synaptic neuron? Chemicals called neurotransmitters are involved in the transmission of impulses at these synapses. The axon terminals contain vesicles filled with these neurotransmitters. When an impulse (action potential) arrives at the axon terminal, it stimulates the movement of the synaptic vesicles towards the membrane where they fuse with the plasma. Figure 18.3 Diagram showing axon terminal and synapse
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# N EURAL CONTROL AND COORDINATION membrane and release their neurotransmitters in the synaptic cleft. The released neurotransmitters bind to their specific receptors, present on the post-synaptic membrane. This binding opens ion channels allowing the entry of ions which can generate a new potential in the post-synaptic neuron. The new potential developed may be either excitatory or inhibitory. # 18.4 C ENTRAL NEURAL SYSTEM The brain is the central information processing organ of our body, and acts as the ‘command and control system’. It controls the voluntary movements, balance of the body, functioning of vital involuntary organs (e.g., lungs, heart, kidneys, etc.), thermoregulation, hunger and thirst, circadian (24-hour) rhythms of our body, activities of several endocrine glands and human behaviour. It is also the site for processing of vision, hearing, speech, memory, intelligence, emotions and thoughts. The human brain is well protected by the skull. Inside the skull, the brain is covered by cranial meninges consisting of an outer layer called dura mater, a very thin middle layer called arachnoid and an inner layer (which is in contact with the brain tissue) called pia mater. The brain can be divided into three major parts: (i) forebrain, (ii) midbrain, and (iii) hindbrain. # Figure 18.4 Diagram showing sagital section of the human brain Cerebral hemisphere
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# BIOLOGY # 18.4.1 Forebrain The forebrain consists of cerebrum, thalamus and hypothalamus (Figure 18.4). Cerebrum forms the major part of the human brain. A deep cleft divides the cerebrum longitudinally into two halves, which are termed as the left and right cerebral hemispheres. The hemispheres are connected by a tract of nerve fibres called corpus callosum. The layer of cells which covers the cerebral hemisphere is called cerebral cortex and is thrown into prominent folds. The cerebral cortex is referred to as the grey matter due to its greyish appearance. The neuron cell bodies are concentrated here giving the colour. The cerebral cortex contains motor areas, sensory areas and large regions that are neither clearly sensory nor motor in function. These regions called as the association areas are responsible for complex functions like intersensory associations, memory and communication. Fibres of the tracts are covered with the myelin sheath, which constitute the inner part of cerebral hemisphere. They give an opaque white appearance to the layer and, hence, is called the white matter. The cerebrum wraps around a structure called thalamus, which is a major coordinating centre for sensory and motor signaling. Another very important part of the brain called hypothalamus lies at the base of the thalamus. The hypothalamus contains a number of centres which control body temperature, urge for eating and drinking. It also contains several groups of neurosecretory cells, which secrete hormones called hypothalamic hormones. The inner parts of cerebral hemispheres and a group of associated deep structures like amygdala, hippocampus, etc., form a complex structure called the limbic lobe or limbic system. Along with the hypothalamus, it is involved in the regulation of sexual behaviour, expression of emotional reactions (e.g., excitement, pleasure, rage and fear), and motivation. # 18.4.2 Midbrain The midbrain is located between the thalamus/hypothalamus of the forebrain and pons of the hindbrain. A canal called the cerebral aqueduct passes through the midbrain. The dorsal portion of the midbrain consists mainly of four round swellings (lobes) called corpora quadrigemina. # 18.4.3 Hindbrain The hindbrain comprises pons, cerebellum and medulla (also called the medulla oblongata). Pons consists of fibre tracts that interconnect different regions of the brain. Cerebellum has very convoluted surface in order to provide the additional space for many more neurons. The medulla of the brain is connected to the spinal cord. The medulla contains centres which control respiration, cardiovascular reflexes and gastric secretions. Three major regions make up the brain stem; mid brain, pons and medulla oblongata. Brain stem forms the connections between the brain and spinal cord.
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# N EURAL CONTROL AND COORDINATION # SUMMARY The neural system coordinates and integrates functions as well as metabolic and homeostatic activities of all the organs. Neurons, the functional units of the neural system, are excitable cells due to a differential concentration gradient of ions across the membrane. The electrical potential difference across the resting neural membrane is called the ‘resting potential’. The nerve impulse is conducted along the axon membrane in the form of a wave of depolarisation and repolarisation. A synapse is formed by the membranes of a pre-synaptic neuron and a post-synaptic neuron which may or may not be separated by a gap called synaptic cleft. Chemicals involved in the transmission of impulses at chemical synapses are called neurotransmitters. Human neural system consists of two parts: (i) central neural system (CNS) and (ii) the peripheral neural system. The CNS consists of the brain and spinal cord. The brain can be divided into three major parts: (i) forebrain, (ii) midbrain, and (iii) hindbrain. The forebrain consists of cerebrum, thalamus, and hypothalamus. The cerebrum is longitudinally divided into two halves that are connected by the corpus callosum. A very important part of the forebrain called hypothalamus controls the body temperature, eating, and drinking. Inner parts of cerebral hemispheres and a group of associated deep structures form a complex structure called limbic system which is concerned with olfaction, autonomic responses, regulation of sexual behaviour, expression of emotional reactions, and motivation. The midbrain receives and integrates visual, tactile, and auditory inputs. The hindbrain comprises pons, cerebellum, and medulla. The cerebellum integrates information received from the semicircular canals of the ear and the auditory system. The medulla contains centres, which control respiration, cardiovascular reflexes, and gastric secretions. Pons consist of fibre tracts that interconnect different regions of the brain. # EXERCISES 1. Briefly describe the structure of the Brain 2. Compare the following: 1. (a) Central neural system (CNS) and Peripheral neural system (PNS) 2. (b) Resting potential and action potential 3. Explain the following processes: 1. (a) Polarisation of the membrane of a nerve fibre 2. (b) Depolarisation of the membrane of a nerve fibre 3. (c) Transmission of a nerve impulse across a chemical synapse 2024-25
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# BIOLOGY 1. Draw labelled diagrams of the following: 1. Neuron 2. Brain 2. Write short notes on the following: 1. Neural coordination 2. Forebrain 3. Midbrain 4. Hindbrain 5. Synapse 3. Give a brief account of Mechanism of synaptic transmission. 4. Explain the role of Na+ in the generation of action potential. 5. Differentiation between: 1. Myelinated and non-myelinated axons 2. Dendrites and axons 3. Thalamus and Hypothalamus 4. Cerebrum and Cerebellum 6. Answer the following: 1. Which part of the human brain is the most developed? 2. Which part of our central neural system acts as a master clock? 7. Distinguish between: 1. afferent neurons and efferent neurons 2. impulse conduction in a myelinated nerve fibre and unmyelinated nerve fibre 3. cranial nerves and spinal nerves. 2024-25
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# Crtolineal # UNIT 3 # CELL: STRUCTURE AND FUNCTIONS # Chapter 8 Biology is the study of living organisms. The detailed description of their form and appearance only brought out their diversity. It is the cell theory that emphasised the unity underlying this diversity of forms, i.e., the cellular organisation of all life forms. A description of cell structure and cell growth by division is given in the chapters comprising this unit. Cell theory also created a sense of mystery around living phenomena, i.e., physiological and behavioural processes. This mystery was the requirement of integrity of cellular organisation for living phenomena to be demonstrated or observed. In studying and understanding the physiological and behavioural processes, one can take a physico-chemical approach and use cell-free systems to investigate. This approach enables us to describe the various processes in molecular terms. The approach is established by analysis of living tissues for elements and compounds. It will tell us what types of organic compounds are present in living organisms. In the next stage, one can ask the question: What are these compounds doing inside a cell? And, in what way they carry out gross physiological processes like digestion, excretion, memory, defense, recognition, etc. In other words we answer the question, what is the molecular basis of all physiological processes? It can also explain the abnormal processes that occur during any diseased condition. This physico-chemical approach to study and understand living organisms is called ‘Reductionist Biology’. The concepts and techniques of physics and chemistry are applied to understand biology. In Chapter 9 of this unit, a brief description of biomolecules is provided. # Chapter 9 # Biomolecules # Chapter 10 # Cell Cycle and Cell Division 2024-25
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# G.N. Ramachandran G.N. Ramachandran, an outstanding figure in the field of protein structure, was the founder of the ‘Madras school’ of conformational analysis of biopolymers. His discovery of the triple helical structure of collagen published in Nature in 1954 and his analysis of the allowed conformations of proteins through the use of the ‘Ramachandran plot’ rank among the most outstanding contributions in structural biology. He was born on October 8, 1922, in a small town, not far from Cochin on the southwestern coast of India. His father was a professor of mathematics at a local college and thus had considerable influence in shaping Ramachandran’s interest in mathematics. After completing his school years, Ramachandran graduated in 1942 as the top-ranking student in the B.Sc. (Honors) Physics course of the University of Madras. He received a Ph.D. from Cambridge University in 1949. While at Cambridge, Ramachandran met Linus Pauling and was deeply influenced by his publications on models of the α-helix and β-sheet structures that directed his attention to solving the structure of collagen. He passed away at the age of 78, on April 7, 2001. (1922 – 2001) 2024-25
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# CHAPTER 8 # CELL: THE UNIT OF LIFE # 8.1 What is a Cell? When you look around, you see both living and non-living things. You must have wondered and asked yourself – ‘what is it that makes an organism living, or what is it that an inanimate thing does not have which a living thing has’? The answer to this is the presence of the basic unit of life – the cell in all living organisms. # 8.2 Cell Theory All organisms are composed of cells. Some are composed of a single cell and are called unicellular organisms while others, like us, composed of many cells, are called multicellular organisms. # 8.3 An Overview of Cell Unicellular organisms are capable of (i) independent existence and (ii) performing the essential functions of life. Anything less than a complete structure of a cell does not ensure independent living. Hence, cell is the fundamental structural and functional unit of all living organisms. # 8.4 Prokaryotic Cells Anton Von Leeuwenhoek first saw and described a live cell. Robert Brown later discovered the nucleus. The invention of the microscope and its improvement leading to the electron microscope revealed all the structural details of the cell. # 8.5 Eukaryotic Cells In 1838, Matthias Schleiden, a German botanist, examined a large number of plants and observed that all plants are composed of different kinds of cells which form the tissues of the plant. At about the same time, Theodore
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# BIOLOGY Schwann (1839), a British Zoologist, studied different types of animal cells and reported that cells had a thin outer layer which is today known as the ‘plasma membrane’. He also concluded, based on his studies on plant tissues, that the presence of cell wall is a unique character of the plant cells. On the basis of this, Schwann proposed the hypothesis that the bodies of animals and plants are composed of cells and products of cells. Schleiden and Schwann together formulated the cell theory. This theory however, did not explain as to how new cells were formed. Rudolf Virchow (1855) first explained that cells divided and new cells are formed from pre-existing cells (Omnis cellula-e cellula). He modified the hypothesis of Schleiden and Schwann to give the cell theory a final shape. Cell theory as understood today is: - (i) all living organisms are composed of cells and products of cells. - (ii) all cells arise from pre-existing cells. # 8.3 A N OVERVIEW OF CELL You have earlier observed cells in an onion peel and/or human cheek cells under the microscope. Let us recollect their structure. The onion cell which is a typical plant cell, has a distinct cell wall as its outer boundary and just within it is the cell membrane. The cells of the human cheek have an outer membrane as the delimiting structure of the cell. Inside each cell is a dense membrane bound structure called nucleus. This nucleus contains the chromosomes which in turn contain the genetic material, DNA. Cells that have membrane bound nuclei are called eukaryotic whereas cells that lack a membrane bound nucleus are prokaryotic. In both prokaryotic and eukaryotic cells, a semi-fluid matrix called cytoplasm occupies the volume of the cell. The cytoplasm is the main arena of cellular activities in both the plant and animal cells. Various chemical reactions occur in it to keep the cell in the ‘living state’. Besides the nucleus, the eukaryotic cells have other membrane bound distinct structures called organelles like the endoplasmic reticulum (ER), the golgi complex, lysosomes, mitochondria, microbodies and vacuoles. The prokaryotic cells lack such membrane bound organelles. Ribosomes are non-membrane bound organelles found in all cells – both eukaryotic as well as prokaryotic. Within the cell, ribosomes are found not only in the cytoplasm but also within the two organelles – chloroplasts (in plants) and mitochondria and on rough ER. Animal cells contain another non-membrane bound organelle called centrosome which helps in cell division. Cells differ greatly in size, shape and activities (Figure 8.1). For example, Mycoplasmas, the smallest cells, are only 0.3 μm in length while bacteria.
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# CELL: THE UNIT OF LIFE # 89 | | | |White blood cells|Columnar epithelial cells| | |---|---|---|---|---|---| |Red blood cells| | |(amoeboid)|(long and narrow)| | |(round and biconcave)|Nerve cell| | | | | | | |(Branched and long)|Mesophyll cells| | | | | | | |A tracheid|(round and oval)| | | | | |(elongated)| | Figure 8.1 Diagram showing different shapes of the cells could be 3 to 5 μm. The largest isolated single cell is the egg of an ostrich. Among multicellular organisms, human red blood cells are about 7.0 μm in diameter. Nerve cells are some of the longest cells. Cells also vary greatly in their shape. They may be disc-like, polygonal, columnar, cuboid, thread like, or even irregular. The shape of the cell may vary with the function they perform. # 8.4 PROKARYOTIC CELLS The prokaryotic cells are represented by bacteria, blue-green algae, mycoplasma and PPLO (Pleuro Pneumonia Like Organisms). They are generally smaller and multiply more rapidly than the eukaryotic cells (Figure 8.2). They may vary greatly in shape and size. The four basic shapes of bacteria are bacillus (rod like), coccus (spherical), vibrio (comma shaped) and spirillum (spiral). The organisation of the prokaryotic cell is fundamentally similar even though prokaryotes exhibit a wide variety of shapes and functions. All 2024-25
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# BIOLOGY Prokaryotes have a cell wall surrounding the cell membrane except in mycoplasma. The fluid matrix filling the cell is the cytoplasm. There is no well-defined nucleus. The genetic material is basically naked, not enveloped by a nuclear membrane. In addition to the genomic DNA (the single chromosome/circular DNA), many bacteria have small circular DNA outside the genomic DNA. These smaller DNA are called plasmids. The plasmid DNA confers certain unique phenotypic characters to such bacteria. One such character is resistance to antibiotics. # Typical bacteria (1-2mm) # PPLO (about 0.1mm) # Viruses (0.02-0.2mm) A typical eukaryotic cell (10-20mm) Figure 8.2: Diagram showing comparison of eukaryotic cell with other eukaryotes # 8.4.1 Cell Envelope and its Modifications Most prokaryotic cells, particularly the bacterial cells, have a chemically complex cell envelope. The cell envelope consists of a tightly bound three layered structure i.e., the outermost glycocalyx followed by the cell wall and then the plasma membrane. Although each layer of the envelope performs distinct function, they act together as a single protective unit. Bacteria can be classified into two groups on the basis of the differences in the cell envelopes and the manner in which they respond to the staining procedure developed by Gram viz., those that take up the gram stain are Gram positive and the others that do not are called Gram negative bacteria. The glycocalyx differs in composition and thickness among different bacteria. It could be a loose sheath called the slime layer in some, while in others it may be thick and tough, called the capsule. The cell wall determines the shape of the cell and provides a strong structural support to prevent the bacterium from bursting or collapsing. The plasma membrane is selectively permeable in nature and interacts with the outside world. This membrane is similar structurally to that of the eukaryotes. A special membranous structure is the mesosome which is formed by the extensions of plasma membrane into the cell. These extensions are in the form of vesicles, tubules and lamellae. They help in cell wall.
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# CELL: THE UNIT OF LIFE formation, DNA replication and distribution to daughter cells. They also help in respiration, secretion processes, to increase the surface area of the plasma membrane and enzymatic content. In some prokaryotes like cyanobacteria, there are other membranous extensions into the cytoplasm called chromatophores which contain pigments. Bacterial cells may be motile or non-motile. If motile, they have thin filamentous extensions from their cell wall called flagella. Bacteria show a range in the number and arrangement of flagella. Bacterial flagellum is composed of three parts – filament, hook and basal body. The filament is the longest portion and extends from the cell surface to the outside. Besides flagella, Pili and Fimbriae are also surface structures of the bacteria but do not play a role in motility. The pili are elongated tubular structures made of a special protein. The fimbriae are small bristle like fibres sprouting out of the cell. In some bacteria, they are known to help attach the bacteria to rocks in streams and also to the host tissues. # 8.4.2 Ribosomes and Inclusion Bodies In prokaryotes, ribosomes are associated with the plasma membrane of the cell. They are about 15 nm by 20 nm in size and are made of two subunits - 50S and 30S units which when present together form 70S prokaryotic ribosomes. Ribosomes are the site of protein synthesis. Several ribosomes may attach to a single mRNA and form a chain called polyribosomes or polysome. The ribosomes of a polysome translate the mRNA into proteins. Inclusion bodies: Reserve material in prokaryotic cells are stored in the cytoplasm in the form of inclusion bodies. These are not bound by any membrane system and lie free in the cytoplasm, e.g., phosphate granules, cyanophycean granules and glycogen granules. Gas vacuoles are found in blue green and purple and green photosynthetic bacteria. # 8.5 EUKARYOTIC CELLS The eukaryotes include all the protists, plants, animals and fungi. In eukaryotic cells there is an extensive compartmentalisation of cytoplasm through the presence of membrane bound organelles. Eukaryotic cells possess an organised nucleus with a nuclear envelope. In addition, eukaryotic cells have a variety of complex locomotory and cytoskeletal structures. Their genetic material is organised into chromosomes. All eukaryotic cells are not identical. Plant and animal cells are different as the former possess cell walls, plastids and a large central vacuole which are absent in animal cells. On the other hand, animal cells have centrioles which are absent in almost all plant cells (Figure 8.3).
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d1f2f59d-ed90-4f32-be58-071e811441ac
# BIOLOGY # Figure 8.3 Diagram showing: # (a) Plant cell |Rough endoplasmic reticulum|Smooth endoplasmic reticulum|Lysosome| |---|---|---| |Plasmodesmata|Nucleus|Nucleolus| |Golgi apparatus|Microtubule|Nuclear envelope| |Plasma membrane|Vacuole|Middle lamella| |Cell wall|(a) Peroxisome|Mitochondrion| |Cytoplasm|Ribosomes|Chloroplast| # (b) Animal cell |Golgi apparatus|Microvilli|Plasma membrane| |---|---|---| |Centriole|Smooth endoplasmic reticulum|Peroxisome| |Lysosome|Ribosomes|Nuclear envelope| |Nucleolus|Mitochondrion|Rough endoplasmic reticulum| |Nucleus|(b)|Cytoplasm| 2024-25
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83344975-74a8-4d8e-b7a4-73fd0c7407fc
# CELL: THE UNIT OF LIFE # 8.5.1 Cell Membrane The detailed structure of the membrane was studied only after the advent of the electron microscope in the 1950s. Meanwhile, chemical studies on the cell membrane, especially in human red blood cells (RBCs), enabled the scientists to deduce the possible structure of plasma membrane. These studies showed that the cell membrane is mainly composed of lipids and proteins. The major lipids are phospholipids that are arranged in a bilayer. Also, the lipids are arranged within the membrane with the polar head towards the outer sides and the hydrophobic tails towards the inner part. This ensures that the nonpolar tail of saturated hydrocarbons is protected from the aqueous environment (Figure 8.4). In addition to phospholipids, the membrane also contains cholesterol. Later, biochemical investigation clearly revealed that the cell membranes also possess protein and carbohydrate. The ratio of protein and lipid varies considerably in different cell types. In human beings, the membrane of the erythrocyte has approximately 52 per cent protein and 40 per cent lipids. Depending on the ease of extraction, membrane proteins can be classified as integral and peripheral. Peripheral proteins lie on the surface of the membrane while the integral proteins are partially or totally buried in the membrane. |Sugar|Peripheral Protein| |---|---| |Phospholipid bilayer|Phospholipid bilayer| |Cholesterol|Integral protein| Figure 8.4 Fluid mosaic model of plasma membrane
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# 8.5 Cell Membrane An improved model of the structure of cell membrane was proposed by Singer and Nicolson (1972) widely accepted as fluid mosaic model (Figure 8.4). According to this, the quasi-fluid nature of lipid enables lateral movement of proteins within the overall bilayer. This ability to move within the membrane is measured as its fluidity. The fluid nature of the membrane is also important from the point of view of functions like cell growth, formation of intercellular junctions, secretion, endocytosis, cell division etc. One of the most important functions of the plasma membrane is the transport of the molecules across it. The membrane is selectively permeable to some molecules present on either side of it. Many molecules can move briefly across the membrane without any requirement of energy and this is called the passive transport. Neutral solutes may move across the membrane by the process of simple diffusion along the concentration gradient, i.e., from higher concentration to the lower. Water may also move across this membrane from higher to lower concentration. Movement of water by diffusion is called osmosis. As the polar molecules cannot pass through the nonpolar lipid bilayer, they require a carrier protein of the membrane to facilitate their transport across the membrane. A few ions or molecules are transported across the membrane against their concentration gradient, i.e., from lower to the higher concentration. Such a transport is an energy dependent process, in which ATP is utilised and is called active transport, e.g., Na+/K Pump. # 8.5.2 Cell Wall As you may recall, a non-living rigid structure called the cell wall forms an outer covering for the plasma membrane of fungi and plants. Cell wall not only gives shape to the cell and protects the cell from mechanical damage and infection, it also helps in cell-to-cell interaction and provides barrier to undesirable macromolecules. Algae have cell wall, made of cellulose, galactans, mannans and minerals like calcium carbonate, while in other plants it consists of cellulose, hemicellulose, pectins and proteins. The cell wall of a young plant cell, the primary wall is capable of growth, which gradually diminishes as the cell matures and the secondary wall is formed on the inner (towards membrane) side of the cell. The middle lamella is a layer mainly of calcium pectate which holds or glues the different neighbouring cells together. The cell wall and middle lamellae may be traversed by plasmodesmata which connect the cytoplasm of neighbouring cells. # 8.5.3 Endomembrane System While each of the membranous organelles is distinct in terms of its
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23d89de0-8cb9-4671-93a5-c0985a1a1546
# CELL: THE UNIT OF LIFE structure and function, many of these are considered together as an endomembrane system because their functions are coordinated. The endomembrane system include endoplasmic reticulum (ER), golgi complex, lysosomes and vacuoles. Since the functions of the mitochondria, chloroplast and peroxisomes are not coordinated with the above components, these are not considered as part of the endomembrane system. # 8.5.3.1 The Endoplasmic Reticulum (ER) Electron microscopic studies of eukaryotic cells reveal the presence of a network or reticulum of tiny tubular structures scattered in the cytoplasm that is called the endoplasmic reticulum (ER) (Figure 8.5). Hence, ER divides the intracellular space into two distinct compartments, i.e., luminal (inside ER) and extra luminal (cytoplasm) compartments. The ER often shows ribosomes attached to their outer surface. The endoplasmic reticulum bearing ribosomes on their surface is called rough endoplasmic reticulum (RER). In the absence of ribosomes they appear smooth and are called smooth endoplasmic reticulum (SER). RER is frequently observed in the cells actively involved in protein synthesis and secretion. They are extensive and continuous with the outer membrane of the nucleus. The smooth endoplasmic reticulum is the major site for synthesis of lipid. In animal cells lipid-like steroidal hormones are synthesised in SER. # 8.5.3.2 Golgi apparatus Camillo Golgi (1898) first observed densely stained reticular structures near the nucleus. These were later named Golgi bodies after him. They consist of many flat, disc-shaped sacs or cisternae of 0.5μm to 1.0μm diameter (Figure 8.6). These are stacked parallel to each other. Varied number of cisternae are present in a Golgi complex. The Golgi cisternae are concentrically arranged near the nucleus with distinct convex cis or the forming.
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833c744f-fa99-461c-b0f7-401f23cd4368
# 96 # BIOLOGY The cis and the trans faces of the organelle are entirely different, but interconnected. The golgi apparatus principally performs the function of packaging materials, to be delivered either to the intra-cellular targets or secreted outside the cell. Materials to be packaged in the form of vesicles from the ER fuse with the cis face of the golgi apparatus and move towards the maturing face. This explains, why the golgi apparatus remains in close association with the endoplasmic reticulum. A number of proteins synthesised by ribosomes on the endoplasmic reticulum are modified in the cisternae of the golgi apparatus before they are released from its trans face. Golgi apparatus is the important site of formation of glycoproteins and glycolipids. # 8.5.3.3 Lysosomes These are membrane bound vesicular structures formed by the process of packaging in the golgi apparatus. The isolated lysosomal vesicles have been found to be very rich in almost all types of hydrolytic enzymes (hydrolases – lipases, proteases, carbohydrases) optimally active at the acidic pH. These enzymes are capable of digesting carbohydrates, proteins, lipids and nucleic acids. # 8.5.3.4 Vacuoles The vacuole is the membrane-bound space found in the cytoplasm. It contains water, sap, excretory product and other materials not useful for the cell. The vacuole is bound by a single membrane called tonoplast. In plant cells the vacuoles can occupy up to 90 per cent of the volume of the cell. In plants, the tonoplast facilitates the transport of a number of ions and other materials against concentration gradients into the vacuole, hence their concentration is significantly higher in the vacuole than in the cytoplasm. In Amoeba the contractile vacuole is important for osmoregulation and excretion. In many cells, as in protists, food vacuoles are formed by engulfing the food particles. # 8.5.4 Mitochondria Mitochondria (sing.: mitochondrion), unless specifically stained, are not easily visible under the microscope. The number of mitochondria per cell is variable depending on the physiological activity of the cells. In terms of shape and size also, considerable degree of variability is observed. Typically it is sausage-shaped or cylindrical having a diameter of 0.2-1.0μm (average 0.5μm) and length 1.0-4.1μm. Each mitochondrion is a double.
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3d8c93d1-b20e-40aa-8769-bdb8b18b7a3b
# CELL: THE UNIT OF LIFE # 8.5.5 Plastids Plastids are found in all plant cells and in euglenoides. These are easily observed under the microscope as they are large. They bear some specific pigments, thus imparting specific colours to the plants. Based on the type of pigments plastids can be classified into chloroplasts, chromoplasts and leucoplasts. The chloroplasts contain chlorophyll and carotenoid pigments which are responsible for trapping light energy essential for photosynthesis. In the chromoplasts fat soluble carotenoid pigments like carotene, xanthophylls and others are present. This gives the part of the plant a yellow, orange or red colour. The leucoplasts are the colourless plastids of varied shapes and sizes with stored nutrients: Amyloplasts store carbohydrates (starch), e.g., potato; elaioplasts store oils and fats whereas # Figure 8.7 Structure of mitochondrion (Longitudinal section) |Outer membrane|Inner membrane|Inter-membrane space| |---|---|---| |Matrix|Matrix|Matrix| |Crista|Crista|Crista| membrane-bound structure with the outer membrane and the inner membrane dividing its lumen distinctly into two aqueous compartments, i.e., the outer compartment and the inner compartment. The inner compartment is filled with a dense homogeneous substance called the matrix. The outer membrane forms the continuous limiting boundary of the organelle. The inner membrane forms a number of infoldings called the cristae (sing.: crista) towards the matrix. The cristae increase the surface area. The two membranes have their own specific enzymes associated with the mitochondrial function. Mitochondria are the sites of aerobic respiration. They produce cellular energy in the form of ATP, hence they are called ‘power houses’ of the cell. The matrix also possesses single circular DNA molecule, a few RNA molecules, ribosomes (70S) and the components required for the synthesis of proteins. The mitochondria divide by fission.
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2c4cbb55-9b44-4e16-963a-66d65a7b4980
# 98 # BIOLOGY The aleuroplasts store proteins. Majority of the chloroplasts of the green plants are found in the mesophyll cells of the leaves. These are lens-shaped, oval, spherical, discoid or even ribbon-like organelles having variable length (5-10μm) and width (2-4μm). Their number varies from 1 per cell of the Chlamydomonas, a green alga to 20-40 per cell in the mesophyll. Like mitochondria, the chloroplasts are also double membrane bound. Of the two, the inner chloroplast membrane is relatively less permeable. The space limited by the inner membrane of the chloroplast is called the stroma. A number of organised flattened membranous sacs called the thylakoids, are present in the stroma (Figure 8.8). Thylakoids are arranged in stacks like the piles of coins called grana (singular: granum) or the intergranal thylakoids. In addition, there are flat membranous tubules called the stroma lamellae connecting the thylakoids of the different grana. The membrane of the thylakoids enclose a space called a lumen. The stroma of the chloroplast contains enzymes required for the synthesis of carbohydrates and proteins. It also contains small, double-stranded circular DNA molecules and ribosomes. Chlorophyll pigments are present in the thylakoids. The ribosomes of the chloroplasts are smaller (70S) than the cytoplasmic ribosomes (80S). # 8.5.6 Ribosomes Ribosomes are the granular structures first observed under the electron microscope as dense particles by George Palade (1953). They are composed of ribonucleic acid (RNA) and proteins and are not surrounded by any membrane. The eukaryotic ribosomes are 80S while the prokaryotic ribosomes are 70S. Each ribosome has two subunits, larger and smaller subunits (Fig 8.9). The two subunits of 80S ribosomes are 60S and 40S while that of 70S ribosomes are 50S and 30S. Here ‘S’ (Svedberg’s Unit) stands for the sedimentation coefficient; it is indirectly a measure of density and size. Both 70S and 80S ribosomes are composed of two subunits. # 8.5.7 Cytoskeleton An elaborate network of filamentous proteinaceous structures consisting of microtubules, microfilaments and intermediate filaments present in the cytoplasm is collectively referred to as the cytoskeleton. The cytoskeleton in a cell are involved in many functions such as mechanical support, motility, maintenance of the shape of the cell. 2024-25
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# C ELL: THE U NIT OF L IFE # 8.5.8 Cilia and Flagella Cilia (sing.: cilium) and flagella (sing.: flagellum) are hair-like outgrowths of the cell membrane. Cilia are small structures which work like oars, causing the movement of either the cell or the surrounding fluid. Flagella are comparatively longer and responsible for cell movement. The prokaryotic bacteria also possess flagella but these are structurally different from that of the eukaryotic flagella. The electron microscopic study of a cilium or the flagellum show that they are covered with plasma membrane. Their core called the axoneme, possesses a number of microtubules running parallel to the long axis. The axoneme usually has nine doublets of radially arranged peripheral microtubules, and a pair of centrally located microtubules. Such an arrangement of axonemal microtubules is referred to as the 9+2 array (Figure 8.10). The central tubules are connected by bridges and is also enclosed by a central sheath, which is connected to one of the tubules of each peripheral doublets by a radial spoke. Thus, there are nine radial spokes. The peripheral doublets are also interconnected by linkers. Both the cilium and flagellum emerge from centriole-like structure called the basal bodies. # 8.5.9 Centrosome and Centrioles Centrosome is an organelle usually containing two cylindrical structures called centrioles. They are surrounded by amorphous pericentriolar materials. Both the centrioles in a centrosome lie perpendicular to each other in which each has an organisation like the cartwheel. They are
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# BIOLOGY made up of nine evenly spaced peripheral fibrils of tubulin protein. Each of the peripheral fibril is a triplet. The adjacent triplets are also linked. The central part of the proximal region of the centriole is also proteinaceous and called the hub, which is connected with tubules of the peripheral triplets by radial spokes made of protein. The centrioles form the basal body of cilia or flagella, and spindle fibres that give rise to spindle apparatus during cell division in animal cells. # 8.5.10 Nucleus Nucleus as a cell organelle was first described by Robert Brown as early as 1831. Later the material of the nucleus stained by the basic dyes was given the name chromatin by Flemming. The interphase nucleus (nucleus of a cell when it is not dividing) has highly extended and elaborate nucleoprotein fibres called chromatin, nuclear matrix and one or more spherical bodies called nucleoli (sing.: nucleolus) (Figure 8.11). |Nucleoplasm|Nucleolus|Nuclear pore|Nuclear membrane| |---|---|---|---| |Figure 8.11 Structure of nucleus|Figure 8.11 Structure of nucleus|Figure 8.11 Structure of nucleus|Figure 8.11 Structure of nucleus| |Electron microscopy has revealed that the nuclear envelope, which consists of two parallel membranes with a space between (10 to 50 nm) called the perinuclear space, forms a barrier between the materials present inside the nucleus and that of the cytoplasm. The outer membrane usually remains continuous with the endoplasmic reticulum and also bears ribosomes on it. At a number of places the nuclear envelope is interrupted by minute pores, which are formed by the fusion of its two membranes. These nuclear pores are the passages through which movement of RNA and protein molecules takes place in both directions between the nucleus and the cytoplasm.|Electron microscopy has revealed that the nuclear envelope, which consists of two parallel membranes with a space between (10 to 50 nm) called the perinuclear space, forms a barrier between the materials present inside the nucleus and that of the cytoplasm. The outer membrane usually remains continuous with the endoplasmic reticulum and also bears ribosomes on it. At a number of places the nuclear envelope is interrupted by minute pores, which are formed by the fusion of its two membranes. These nuclear pores are the passages through which movement of RNA and protein molecules takes place in both directions between the nucleus and the cytoplasm.|Electron microscopy has revealed that the nuclear envelope, which consists of two parallel membranes with a space between (10 to 50 nm) called the perinuclear space, forms a barrier between the materials present inside the nucleus and that of the cytoplasm. The outer membrane usually remains continuous with the endoplasmic reticulum and also bears ribosomes on it. At a number of places the nuclear envelope is interrupted by minute pores, which are formed by the fusion of its two membranes. These nuclear pores are the passages through which movement of RNA and protein molecules takes place in both directions between the nucleus and the cytoplasm.|Electron microscopy has revealed that the nuclear envelope, which consists of two parallel membranes with a space between (10 to 50 nm) called the perinuclear space, forms a barrier between the materials present inside the nucleus and that of the cytoplasm. The outer membrane usually remains continuous with the endoplasmic reticulum and also bears ribosomes on it. At a number of places the nuclear envelope is interrupted by minute pores, which are formed by the fusion of its two membranes. These nuclear pores are the passages through which movement of RNA and protein molecules takes place in both directions between the nucleus and the cytoplasm.| Normally, there is only one nucleus per cell, variations in the number of nuclei are also frequently observed. Can you recollect names of organisms that have more than one nucleus per cell? Some mature cells even lack nucleus, e.g., erythrocytes of many mammals and sieve tube cells of vascular plants. Would you consider these cells as ‘living’? The nuclear matrix or the nucleoplasm contains nucleolus and chromatin. The nucleoli are spherical structures present in the nucleoplasm. The content of nucleolus is continuous with the rest of the nucleoplasm as it is not a membrane bound structure. It is a site for active ribosomal RNA synthesis. Larger and more numerous nucleoli are present in cells actively carrying out protein synthesis. 2024-25
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ab2247fc-5637-4fed-af35-33f29988a417
# CELL: THE UNIT OF LIFE You may recall that the interphase nucleus has a loose and indistinct network of nucleoprotein fibres called chromatin. But during different stages of cell division, cells show structured chromosomes in place of the nucleus. Chromatin contains DNA and some basic proteins called histones, some non-histone proteins and also RNA. A single human cell has approximately two metre long thread of DNA distributed among its forty six (twenty three pairs) chromosomes. You will study the details of DNA packaging in the form of a chromosome in class XII. Every chromosome (visible only in dividing cells) essentially has a primary constriction or the centromere on the sides of which disc shaped structures called kinetochores are present (Figure 8.12). Centromere holds two chromatids of a chromosome. Based on the position of the centromere, the chromosomes can be classified into four types (Figure 8.13). The metacentric chromosome has middle centromere forming two equal arms of the chromosome. The sub-metacentric chromosome has centromere slightly away from the middle of the chromosome resulting into one shorter arm and one longer arm. In case of acrocentric chromosome the centromere is situated close to its end forming one extremely short and one very long arm, whereas the telocentric chromosome has a terminal centromere. # Kinetochore Figure 8.12 Chromosome with kinetochore Figure 8.13 Types of chromosomes based on the position of centromere
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67f5bfc1-995e-494f-9e32-4801d555539c
# BIOLOGY Sometimes a few chromosomes have non-staining secondary constrictions at a constant location. This gives the appearance of a small fragment called the satellite. # 8.5.11 Microbodies Many membrane bound minute vesicles called microbodies that contain various enzymes, are present in both plant and animal cells. # SUMMARY All organisms are made of cells or aggregates of cells. Cells vary in their shape, size and activities/functions. Based on the presence or absence of a membrane bound nucleus and other organelles, cells and hence organisms can be named as eukaryotic or prokaryotic. A typical eukaryotic cell consists of a cell membrane, nucleus and cytoplasm. Plant cells have a cell wall outside the cell membrane. The plasma membrane is selectively permeable and facilitates transport of several molecules. The endomembrane system includes ER, golgi complex, lysosomes and vacuoles. All the cell organelles perform different but specific functions. Centrosome and centriole form the basal body of cilia and flagella that facilitate locomotion. In animal cells, centrioles also form spindle apparatus during cell division. Nucleus contains nucleoli and chromatin network. It not only controls the activities of organelles but also plays a major role in heredity. Endoplasmic reticulum contains tubules or cisternae. They are of two types: rough and smooth. ER helps in the transport of substances, synthesis of proteins, lipoproteins and glycogen. The golgi body is a membranous organelle composed of flattened sacs. The secretions of cells are packed in them and transported from the cell. Lysosomes are single membrane structures containing enzymes for digestion of all types of macromolecules. Ribosomes are involved in protein synthesis. These occur freely in the cytoplasm or are associated with ER. Mitochondria help in oxidative phosphorylation and generation of adenosine triphosphate. They are bound by double membrane; the outer membrane is smooth and inner one folds into several cristae. Plastids are pigment containing organelles found in plant cells only. In plant cells, chloroplasts are responsible for trapping light energy essential for photosynthesis. The grana, in the plastid, is the site of light reactions and the stroma of dark reactions. The green coloured plastids are chloroplasts, which contain chlorophyll, whereas the other coloured plastids are chromoplasts, which may contain pigments like carotene and xanthophyll. The nucleus is enclosed by nuclear envelope, a double membrane structure with nuclear pores. The inner membrane encloses the nucleoplasm and the chromatin material. Thus, cell is the structural and functional unit of life. 2024-25
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7c464bb6-3a61-4b6d-971f-c16c281817af
# CELL: THE UNIT OF LIFE # EXERCISES 1. Which of the following is not correct? - (a) Robert Brown discovered the cell. - (b) Schleiden and Schwann formulated the cell theory. - (c) Virchow explained that cells are formed from pre-existing cells. - (d) A unicellular organism carries out its life activities within a single cell. 2. New cells generate from - (a) bacterial fermentation - (b) regeneration of old cells - (c) pre-existing cells - (d) abiotic materials 3. Match the following |Column I|Column II| |---|---| |(a) Cristae|(i)| |(b) Cisternae|(ii) Infoldings in mitochondria| |(c) Thylakoids|(iii)| 4. Which of the following is correct: - (a) Cells of all living organisms have a nucleus. - (b) Both animal and plant cells have a well defined cell wall. - (c) In prokaryotes, there are no membrane bound organelles. - (d) Cells are formed de novo from abiotic materials. 5. What is a mesosome in a prokaryotic cell? Mention the functions that it performs. 6. How do neutral solutes move across the plasma membrane? Can the polar molecules also move across it in the same way? If not, then how are these transported across the membrane? 7. Name two cell-organelles that are double membrane bound. What are the characteristics of these two organelles? State their functions and draw labelled diagrams of both. 8. What are the characteristics of prokaryotic cells? 9. Multicellular organisms have division of labour. Explain. 10. Cell is the basic unit of life. Discuss in brief. 11. What are nuclear pores? State their function. 12. Both lysosomes and vacuoles are endomembrane structures, yet they differ in terms of their functions. Comment. 13. Describe the structure of the following with the help of labelled diagrams. - (i) Nucleus - (ii) Centrosome 14. What is a centromere? How does the position of centromere form the basis of classification of chromosomes. Support your answer with a diagram showing the position of centromere on different types of chromosomes. Flat membranous sacs in stroma Disc-shaped sacs in Golgi apparatus
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