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The structure of the text of this module is presented in the form
of the following flow diagram:

Discovery
and history
Nomenclature
and Glossary
classification

Origin and References
evolution

Structure and
composition Quiz
Module
Structure
Isolation
and Assignments
purification

Replication
FAQs
and
transmission

Economic Nobel laureates
importance in virology

A historical overview of viruses

Viruses have been infecting humans as well as other
organisms for thousands of years, thereby causing millions of
deaths and inflicting huge impact on the course of human history.
These viral outbreaks and the costs associated with them have
made man conscious about them from the very early times.
However, history of the science of virology is not too ancient,
mainly because of lack of tools and techniques to deal with these
infitismally small agents untill recent past. The course of
development of this field of microbiological science witnessed
some abrupt changes. In the dynamic and rapidly progressing
field of contemporary virology, today’s discovery is tomorrow’s
history, and it is beyond the scope of present module to present a
complete history of this discipline. While some of the pioneer and
landmark contributions are listed in this brief historical account of
viruses, many others may not find their place herein.

Discovery of viruses can not be attributed to a single person,
but to the joint contribution of a team of Dutch, German and
Russian scientists. After Louis Pasteur and Robert Koch
discovered that infectious diseases were caused by minute living
organisms or 'germs', it was expected that the germs for all
infectious diseases would be discovered. However, bacteriological
techniques failed to demonstrate the causative organisms for

While Schelsinger (1933) first determined the composition of a virus and showed that a . and demonstrated that the crystals retain their infectivity when inoculated into the healthy plants. It was the Russian pathologist Ivanovsky who in 1892 used the filter devised by Charles Chamberland in 1884 to trap smallest possible bacteria to demonstrate that the juice from tobacco plants infected with the 'mosaic' disease contain disease causing agent smaller than any known bacterium. But in 1935 Stanley purified and crystallized the virus causing tobacco mosaic disease. which is as yet unsettled for many practical purposes. while noticing that some agent was destroying his cultures of bacilli. Bacteriophages (viruses that parasitise bacteria) were discovered by the French scientist d'Herelle (1917). He thus showed that viruses were not like typical cells thereby openening a debate on living and non living nature of viruses. Stanley for his unprecedented discovery was duly rewarded with Noble Prize in Chemistry in 1946. but untill 1930 viruses were considered as extremely small bacteria or even enzymes or proteins. small pox. This mosaic causing agent was later called a virus. though he still considered it a bacterium. rabies and mumps. Beijerinck (1851-1931) deserves the credit for first realizing that contagions of tobacco mosaic differed essentially from micro-organisms.many diseases like measles. Some people argue that viruses are neither totally inert nor completely alive and may better be regarded as active (when multiplying) or inactive (not multiplying).

absence of fossil records and lack of proper knowledge of their origin and evolutionary history. (ii) infection is the result of penetration of viral DNA into cells. mainly on the basis of their nucleic acid type and strandedness and presence or absence of the envelope. Earlier. viruses were generally classified by the organisms they infect. such as animal viruses. plant viruses. While a flawless and widely acceptable classification system of viruses is yet to emerge. or bacteriophages. which was established in 1966 and meets after every four years. Classification of viruses The classification of viruses is more or less problematic and relatively less satisfactory. arranged by . The ICTV. published its seventh report in October 2000. The pioneering contributions by these microbiologists guided the way forward for the recent advances in virology. presence of both living and non-living features. Hershey and Chase (1952) studied the T2 bacteriophage and demonstrated that (i) the genetic information is carried in the phage DNA. mainly due to their ultramicroscopic size.bacteriophage consists of only protein and DNA. recently the International Committee on Taxonomy of Viruses (ICTV) attempted to develop a uniform classification system and divided viruses into 73 families. which is considered as the official reference for virus taxonomy and nomenclature and covers more than 4000 viruses.

The important taxonomic criteria for classification of viruses include: a. f. and j. The ICTV considers ‘Family’ as the highest taxonomic category and accepts Van Regenmortel’s (1990) definition of ‘virus species’ as a polythetic class of viruses that constitutes a replicating lineage and occupies a particular ecological niche. d. or . Intracellular location of the viral replication: cytoplasm or nucleus. Number of capsomers in icosahedral viruses. Capsid symmetry: Icosahedral. c. Immunological properties. i. helical. segmentation and number of pieces or segments. g. bacteria. The abovementioned criteria can be categorized into: i. Mode of transmission. b. The type of disease caused. Diameter of the virion or the nucleocapsid. plant. complex or binal. Presence or absence of the envelope and its sensitivity. structure.. fungi. site of replication). single or double strandedness. etc. molecular weight. h. Nucleic acid characteristics: DNA or RNA. e. Nature of the host: animal. Primary characteristics (such as chemical nature.families.

antegenecity etc. Besides. might represent the direct descendents of the self-replicating units (in primordial soup) from which the first cells evolved and then coevolved with more complex life forms. Notwithstanding the debate on the origin of viruses. also known as Retrogressive evolution. While all these hypotheses (Fig. a unifying concept of their origin is yet to emerge. Secondary characteristics (such as host range. viruses act as agents of evolution by their participation in lateral gene transfer. by multiple origins. ii.1) contribute partly to our uderstanding of viral origin. their rapid evolutionary potential stems from their key attributes including: . because of simplicity of their genome.  Viruses are escaped nucleic acids no longer under control of the cell. the following three major theories have been advanced to explain their origin and evolution:  Viruses.) Origin and evolution of viruses While the origin of viruses is still an intriguing question. mode of transmission. This is also known as the Escaped Gene Theory.  Viruses evolved from the free living organisms that invaded other life forms and gradually lost functions. It is believed that viruses have arisen and probably continue to arise.

1 : An integrated framework for the possible origin and evolution of viruses. • Extremely small genome size • Enormous population sizes • Short generation times. and • High mutation rates. cytoplasm with ribosomes and enzymes necessary for protein synthesis and energy production. Retrogressive Direct evolution due to descendence loss of functions from primitive upon invasion self-replicating of other life units forms Escaped Gene Theory (Escaped nucleic acids no longer controlled by cells) Fig. Structure and composition of viruses Viruses lack cellular organization and do not possess plasma membranes. Viruses range in size .

Viral Genome Viral genome. unlike genomes of rest of the life forms. on the higher side they are of the size of of the smallest bacteria. such as Chlamydeae and Mycoplasmas.g. In fact. All viruses consist of two two basic components: i.from as large as 250 nm (e. on the lower side they have the same diameter as a DNA molecule. However. Parvoviruses). The nucleic acids may be double-stranded or single-stranded. viruses are exceptionally flexible with respect to the nature of their genetic matrerial.the capsid.g.the genome. Genome and capsid together comprise the nucleocapsid or genocapsid. but never both. many viruses possess a typical lipoprotenaceous bimembrane outside their capsids and are known as enveloped viruses. As in cells. Poxviruses) to as small as 20 nm (e. contains either DNA or RNA. A surrounding coat of protein. and employ all the four possible nucleic acid types: . and ii. the nucleic acid of each virus encodes the genetic information for the synthesis of important viral proteins. A core of nucleic acid. Thus.

M13 bacteriophages ii. Single stranded DNA (ssDNA). however. In shape also the viral .g. e. There are two types of RNA-based viruses. Double stranded DNA (dsDNA). e. Brome mosaic virus iv. which must first catalyze the production of complementary messenger RNA from the virion genomic RNA. Parvoviruses. While the smallest genome of the order of 1X106 Daltons (e. the genomic RNA is termed a plus (+) strand because it acts as messenger RNA for direct synthesis (translation) of viral protein. but also the size of genome. Single stranded RNA (ss RNA).0 to 1. have DNA as their genetic material. all the above four types of nucleic acid types are found in animal viruses. have negative (-) of RNA. in viruses varies greatly.g. e. i.g. Lambda phages iii.6 X 108 Dalton (e. TMV. called RNA-dependent RNA polymerase (transcriptase). three types of plant viruses. Double stranded RNA (ds RNA). In most. the virion has an enzyme. Retroviruses. A few.g. Not only the type. Herpesvirus and Vaccinia virus) is large enough to direct the synthesis of over 100 proteins.g. Badnaviruses and Caulimoviruses. While plant viruses are largely ssRNA viruses. before viral protein synthesis can occur. However.g. the largest genomes of the size of 1. In these cases. MS2 and QB viruses) can code for just 3 to 4 protiens only. namely Geminiviruses. e.

This shell is composed of protein organized in subunits known as capsomers. These molecular comparisons have shown considerable similraity in viruses infecting apparently different hosts. and • Facilitation of attachment and penetration of viruses into the host. viral RNA in a liquid suspension of protein molecules will self-assemble a capsid to become a functional and infectious virus. Capsid Capsid is a proteinaceous coat. The important functions of the capsids include: • Protection of the viral genome. The number of capsomers is species specific and is used as a taxonomic character.nucleic acids may vary from linear through circular to segmented or fragmented. or shell-like covering. . It is closely associated with the nucleic acid and reflect its configuration. • Determining shape of the viruses. either a rod-shaped helix or a polygon-shaped sphere. Due to the advances in molecular biology techniques. Under the appropriate conditions. the entire nucleic acid sequences have recently been determined in hundreds of viruses. of nucleic acids in viruses.

. Surrounding nucleocapsids. These viral coded proteins project as spikes. envelope protiens are usually coded by viral genes. Viruses lacking such envelopes are referred to as naked or non-enveloped. Enveloped viruses. many viruses have a typical bimembrane. While the lipids and carbohydrates are normal host constituants. damaged easily in comparison to naked viruses.Envelope Envelope is a membraneous covering found on some virus capsids that allow the virus to enter a host cell by fusing with its cell membrane. The envelope is predominatly acquired from the host cells during the replication process and is unique to each type of viruses. because the membranes are destroyed easily by the harsh environmental factors. The main adventages of having envelopes in viruses include: i.2). Facilitation of infecting new cells through membrane fusion with host cell membrane. in combination with glycogen moities. known as peplomers. determine viral host specificity. and are referred to as enveloped (Fig. ii. Prevention from attack by host’s immune system because envelopes are acquired from and are therefore similar to host cell membranes. such as increased temperatures. which. composed of glycolipids and protiens.

Helical. ii. or other chemical disinfectants. the advent and use of electron microscopy. Fig.freezing and thawing. X- ray differaction. 2. Icosahedral. HIV virus with its lipid envelope Morphological types of viruses While the ultrastructural details of viruses remained unclear for a long period of time after their discovery. biochemical analysis and immunological techniques have provided useful insights into their structure and composition. Viruses are broadly classified into the following morphological types: i. iii. mainly due to lack of proper techniques. Enveloped and . highly acidic or alkaline conditions.

The capsomers in the capsids of icosahedral viruses are composed of protomers with 5 (pentamers) or 6 (hexamers) subunits. While pentamers are found on the vertices of the icosahedron.iv. efficient ways of enclosing and encoding maximum information in minimum space of this small genome is inevitable. The sphere shaped virus is actually a 20-sided polygon (icosahedron). Complex. Icosahedral In view of the exceptionally small genome size of viruses. hexamers form the edges of its triangular faces. The examples of icosahedral viruses include Adenoviruses. which has only pentamers. Most icosahedral capsids apparently contain both pentamers and hexamers. a small dsDNA virus. . Icosahedron is one of the nature’s most favourite shapes to accomplish this in case of viruses. except in Simian virus 40 (SV 40). A typical icosahedron is a polyhedron with 12 vertices in 20 equilateral triangles as faces (Fig. 3). Polioviruses and Canine Parvoviruses.

3. They may be very rigid (e. TMV) or flexible (e. In such viruses.g. TMV. Influenza virus) depending upon the nature of the constituant proteins.g. Fig. the protein subunits are arranged in helical spiral with genome fitted into a groove on the inner portion of the protein. The size of the helical capsid is influenced by both its protomers and the nucleic acid enclosed within the capsid.g. e. long. The helical viruses are narrow (15-20nm) and may be quite long (300-400nm). . Icosahedron Helical Helical viruses are the rod shaped. hollow cylinders with linear array of the nucleic acid and the protein subunits making up the capsid.

Complex bacterial viruses with both head and tail are said to have binal symmetry. because they possess a combination of icosahedral (the head) and helical (the tail) symmetry.4. and do fit neither in icosahedral nor in helical capsid symmetry. The Poxviruses and large bacteriophages are two important examples. tail. Helical virus Complex viruses These are the viruses that are assembed from parts that are synthesised separately (head. capsomer). but have complicated morphology. Fig. .

Viral replication Viral replication refers to the means by which virus particles make new copies of themselves. The newly formed virions may leave the host cell either without causing any damage to the host cells (Lysogenic cycle) or sometimes killing them in the process (Lytic cycle). . Viruses replicate by infecting the host cells and overtaking their synthetic capabilities to produce more viruses.

Fig.6. plants and bacteria exhibit essentially similar sequence of major events in the process of replication. Lytic and lysogenic cycles of viral replication. which generally takes place through rendom collision of virion . Adsorption Adsorption is the first step in viral replication. Viral replication can be principally divided into several stages as follows: 1. viruses of animals. Although precise mechanisms of viral replication may vary.

However. 2. through endosomes at the cell surface. in animal viruses the receptor sites are distributed over th entire capsid surface. Bacteriophages and animal viruses have attachment sites facilitating them to attach to complementry receptor sites on host cell surfaces. unlike bacteriophages which have tails as attachment sites. Penetration and uncoating Depending upon presence or absence of the envelope. The viral attachment protein recognizes specific receptors. viruses enter the cell in a variety of ways according to their nature. carbohydrate or lipid. or ii. Distribution of specific receptor proteins on the host cell membrane plays a crucial role in dtermining the host-specificity of viruses. Attachment is via ionic interactions. Some enveloped viruses fuse directly with the plasma membrane. Cells without appropriate receptors are not susceptible to viruses. which may be protein. In enveloped viruses entry may be: i.particles with host cell surfaces. Thus. which are temperature-independent. by fusing with the plasma membrane. . outside of the cell. the internal components of the virion are immediately delivered to the cytoplasm of the cell.

Uncoating The process in which protein coats of animal viruses that have entered cells. As the endosomes become acidified. Synthesis of viral nucleic acids and protein Many strategies are used by viruses for synthesising nucleic acids and proteins. When the nucleic acid is uncoated. Nucleic acid has to be sufficiently uncoated so that virus replication can begin at this stage. the latent fusion activity of the virus proteins becomes activated by the fall in pH and the virion membrane fuses with the endosome membrane. They then cross (or destroy) the endosomal membrane. infectious virus particles cannot be recovered from the cell. There is an ECLIPSE phase which lasts until new infectious virions are made. are removed by proteolytic enzymes. The non-enveloped viruses may cross the plasma membrane directly or may be taken up into endosomes. 3. is called uncoating. Some enveloped viruses require an acid pH for fusion to occur and are unable to fuse directly with the plasma membrane. These viruses are taken up by invagination of the membrane into endosomes. This results in delivery of the internal components of the virus to the cytoplasm of the cell. and many variations occur between .

if enveloped. RNA viruses replicate their nucleic acid by a greater variety of ways than DNA viruses on the basis of which they have been categorized into four classes. There may be a maturation step that follows the initial assembly process. constitutes maturartion. 5. Pox viruses are an exception. may bud from the cell. where it takes place in the cytoplasm. some budding viruses may . 4. Release Mechanism of virus release differ between naked and enveloped vruses. Assembly/maturation Once abundant viral genomes. or.even if they make no contribution to the morphology or rigidity of the virion .non-structural proteins are those viral proteins found in the cell but not packaged into the virion. in which new virus particles are assembled. All proteins in a mature virus particle are said to be structural proteins . Thus. Budding viruses do not necessarily kill the cell. assembly of these components in the complete virions starts. While DNA replication usually occurs in the host cell nucleus.DNA viruses and RNA viruses. This process. enzymes and other proteins have been synthesised. Virus may be released due to cell lysis. A detailed description of these mechanisms is beyond the scope of present module.

ii. Since viruses are inactive outside their hosts. Most common ways and means of viral transmission in plants include: i. Isolation and purification of viruses Isolation of viruses from their hosts and subsequent purification assume pivotal significance for understanding their physico-chemical nature. Transmission of viruses Transmission refers to the transfer of viruses from one host to the other through different carriers or agencies. Mechanical transmission (natural or artificial). etc. Through plant propagules. Through soil inhabiting organisms. Not all released viral particles are infectious. carriers or plants or indirect contact with inanimate objects such as contaminated food and water. iii. With dodder. Via insects and mites. The ratio of non-infectious to infectious particles varies with the virus and the growth conditions. By grafting. be able to set up persistent infections. iv. composition. and vi. they must undergo passive transfer from one plant to the other. replication and mode of . They transmit either by direct contact with infected individuals. v.

More stability than normal host cell components. Chromatography After successful isolation. Precipitation c.penetration. Filtration f. Various techniques employed for virus isolation and purification include: a. Denaturation d. depend on several viral characteristics. Purification usually requires repetition of above techniques at different intensities for different periods. Electrophoresis g. Presence of surface proteins. the objective of purification is high yield of viruses with lowest amount of contaminants and with highest degree of purity or integrity and infectivity. Centrifugation (differential and density gradient) b. such as: i. however. ii. Isolation techniques. Larger size of viruses relative to proteins.000 kinds of plant diseases are known to be caused by more than 600 different kinds of identified plant . and iii. Enzymatic digestion e. Economic importance of viruses Well over 2.

Losses incured by viruses may be: a. and buffalo. Disease incidence. and iii. Beneficial aspects of viruses . vitality. influenza and AIDS. Most common human diseases caused by viruses include pox. foot and mouth disease of cow. ii. The commonly occuring viral diseases in cattle iclude. polio. more than 400 different viruses can infect humans. except in a few cases. sheep. Viral diseases greatly reduce the productivity of many kinds of agricultural and horticultural crops. Three parameters that may be considered as quantitative indicators for assesment of losses include: i. thereby causing myriad of diseases with no or just prophylactic or supportive treatment. Biological (Damage inflicted upon the organism) b. hum. goat. Worldwide losses due to viral diseases are estimated at about $15 billion annually. quality and the market value of the organisms infected.viruses. Duration of infection. Besides. Economic (What finally counts through reduced production) The losses are estimated in terms of reduction in growth. Disease severity.

especially ornamentals. While viruses are generally considered harmful for the reasons stated above.They are said to cause many plant diseases and are also referred to as subviral pathogens. Thus. vaccines are produced from viruses.O. such as: i) Different vaccines. Their discovery was made by a team of scientists led by T. Subsequently it was established that RNA is the causative agent for PSTV and currently more than a dozen of plant diseases are known to be caused by viroids. ii) Viruses comprise the significant experimental models and tools for scientific studies in molecular biology and genetic engineering. caused by viruses has aesthetic appeal.Diener (1971) of USDA who found that nothing destroyed the Potato Spindle Tuber Disease Causing Agent (PSTV) except the RNAase. yet some viruses can play positive role in human welfare. Viroids cause plant diseases by interfering with mRNA processing. jaundice etc. like small pox. Viroids Infectious RNA molecules that function independently without encapsidation by any protein coat are called as viroids. the RNA degrading enzymes. . iii) Variegated foliage of some plants. viroids are very different from viruses and each viroid is solely a small RNA molecule.

Research indicates that prions are normal proteins that become folded incorrectly. Prions Proteinaceous infectious particles are called as prions. including Creutzfeldt-Jakob disease. kuru. investigations of prion diseases suggest that new strategies for the prevention and treatment of these disorders may also find application in the more common degenerative diseases. Prions cause neurological degenerative diseases. Virusoids are similar to viroids in . and it only encodes structural proteins. The genome of virusoids consist of several hundred nucleotides. the best studied prions are the ones that cause degenerative disorders of the central nervous system (called as scrapie) in sheep and goat. While knowledge about prions has profound implications for studies of the structural plasticity of proteins. but not to the nucleases. and not the otherway round. Prions provide the only example were genetic information flows from proteins to nucleic acids. prions are suceptible to protein-digesting enzymes. Virusoids Virusoids are circular single-stranded RNAs dependent on plant viruses for replication and encapsidation. scrapie and Mad Cow disease by an entirely novel mechanism. However. Named by Stanley Prusiner.

" a separate nucleic acid not part of the viral chromosome. they are classified as satellites. while being studied in virology. An example of a "helper" virus is the subterranean clover mottle virus. are not considered as viruses but as subviral particles. Virusoid genomes are 220 to 388 nucleotides long and do not code for any proteins. Virus enzymes may aid replication of the virusoid RNA. The virusoid is incorporated into the virus particle and transmitted as a "satellite. which has an associated virusoid. but each requires that the cell be infected with a specific "helper" virus. . and the helper viruses for these are all members of the Sobemovirus family. They. structure and means of replication (rolling-circle replication). Virusoids can replicate in the cytoplasm and possess ribozyme activity. Five virusoids are known. RNA replication is similar to that of viroids. Replication of the helper virus is independent of the virusoid. The term virusoid is also sometimes used more generally to refer to all the satellites. but instead serve only to replicate themselves.size. Since they depend on helper viruses.

Mechanism of virus Harshey & Luria infection in living cells 1975 Baltimore. Interaction between Temin & tumour viruses and Dulbecco genetic material of . Weller Cultivation of poli & Robbins virus 1966 Rous Viruses and cancer 1969 Delbruck. Noble prizes awarded in virology Year Prize Topic Studied of Winner(s) Prize 1946 Stainley Crystallization of viruses 1951 Theiler Vaccine for yellow fever 1954 Enders.

the cell 1996 Doherry & Recognition of virus Zinkernagel infected cells by immune defenses 1997 Stanley Prions Prusiner 2008 Human papilloma Harald zur viruses causing Hausen cervical cancer Françoise Human Barré-Sinoussi immunodeficiency and Luc virus Montagnier .