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).

Let us now look at individual cell organelles to understand their structure and functions.

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 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 membrane while the integral proteins are partially or totally buried in the 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.

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.

Endomembrane System

While each of the membranous organelles is distinct in terms of its 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.

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 reticulun 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.

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 \(0.5 \mu m\) to \(1.0 \mu m\) 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 face and concave trans or the maturing face. 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.

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.

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.

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 \(0.2-1.0 \mu m\) (average \(0.5 \mu m\) ) and length \(1.0-4.1 \mu m\). Each mitochondrion is a double 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 (Figure 8.7). 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.

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 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 ( \(5-10 \mu m\) ) and width \((2-4 \mu m)\). 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, doublestranded 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).

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.

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.

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.

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 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.

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). 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.

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.

Sometimes a few chromosomes have non-staining secondary constrictions at a constant location. This gives the appearance of a small fragment called the satellite.

Microbodies

Many membrane bound minute vesicles called microbodies that contain various enzymes, are present in both plant and animal cells.