Drugs

Antibacterial agents (antibiotics)

Antibacterial antibiotics act at a variety of sites. However, in many cases, they act by either making the plasma membrane of bacteria more permeable to essential ions and other small molecules by ionophoric action or by inhibiting cell wall synthesis. Those compounds that act on the plasma membrane also have the ability to penetrate the cell wall structure. In both cases, the net result is a loss in the integrity of the fungal cell envelope, which leads to irreversible cell damage and death.

 

Ionophoric antibiotic action

Ionophores are substances that can penetrate a cell membrane and increase its permeability

to ions. They may be naturally occurring compounds such as the antibiotic gramicidin A produced by Bacillus brevis and valinomycin obtained from Streptomyces fulvissimus, or synthetic compounds like the crown and cryptate compounds (Fig.). However, ionophores transport ions in both directions across a membrane. Consequently, they will only reduce the concentration of a specific ion until its concentration is the same on both sides of a membrane. However, a number of drugs are believed to owe their action to the ionophoric transfer of essential substances out of the cell.

 

Examples of naturally occurring and synthetic ionophores

 

The general mode of action of ionophores in ion transport. (a) A channel formed by two gramicidin A molecules, N-terminal to N-terminal. (b) The sequence of events in the operation of a carrier ionophore such as valinomycin

Ionophores operate in two ways:

1. They form channels across the membrane through which ions can diffuse down a concentration gradient (Fig.a).

2. They act as carriers that pick up the ion on one side of the membrane, transport it across, and release it into the fluid on the other side of the membrane (Fig. b).

The structure of each channel in a Channel ionophore is characteristic of the channel-former. For example, gramicidin forms a channel (tube) composed of two molecules whose N-terminals meet in the middle of the membrane.

 

Each gramicidine molecule is in the form of a left-handed helix, which results in the polar groups lining the interior of the channel. This facilitates the transfer of polar ions through the channel. A single gramicidin channel can allow the transport of up to 107 K+ ions per second.

Carrier ionophores are specific for particular ions. For example, valinomycin will transport K+ ions but not Na2+ or Li+ ions. It is believed to form an octahedral complex with six carbonyl group oxygen atoms acting as ligands. The resulting chelation complex has a hydrophobic exterior that allows the complex to diffuse through the membrane. However, the rigid nature of the molecule coupled with its size makes the binding site of valinomycin too large to form octahedral complexes with the smaller Na2+and Li+ ions. Consequently, it is more energetically favourable for Na2+ and Li+ ions to remain in solution as their hydrated ions. Ionophores are mainly active against Gram-positive bacteria. However, until now, most of the compounds examined do not significantly differentiate between bacterial and mammalian membranes and so are of little clinical use. However, they are of considerable use as research tools in the investigation of drug action.

 

Cell wall synthesis inhibition

The cell walls of all bacteria are being continuously replaced because they are continuously being broken down by enzymes in the extracellular fluid. Antibacterial agents can inhibit this replacement biosynthesis of the cell wall at any stage in its formation. Investigations using Staphylococcus aureus have yielded a great deal of detail about the biosynthesis of its cell wall but there are still areas of the biosynthesis that have not yet been fully elucidated. Experimental investigations of the cell wall synthesis of other bacterial species suggest that similar routes are followed. A detailed knowledge of the route followed and the enzymes involved is an extremely useful prerequisite in the quest for new drug substances.

 

It is convenient to introduce the subject of antibacterial action due to inhibition of cell wall synthesis by dividing the synthesis into three stages:

1. The formation of precursor starting materials.

2. The formation of the peptidoglycan chains.

3. The cross-linking of the peptidoglycan chains.

However, it should be realised that not only can an antibiotic inhibit cell wall formation but it may also have other areas of action such as the plasma membrane of a bacterium. Furthermore, it is emphasised that the biochemical pathways discussed are a simplification based on experimental evidence. However, it is likely that the drugs act in the same manner

on other susceptible bacteria.

Drugs that inhibit the formation of the starting compounds A convenient starting point for cell wall synthesis is N-acetylglucosamine-1-phosphate (NAG-1-P), which is found in all life forms. This compound is believed to react with uridine triphosphate (UTP) to form uridine diphospho-N-acetylglucosamine (UDPNAG), one of the precursors of the peptidoglycan chain (Fig.). Some of the UDPNAG is further converted by a series of steps into the uridine diphospho-N acetylmuramic acid pentapeptide derivative (UDPNAM-pentapeptide), the second precursor of the peptidoglycan polymer chain. Drug action can inhibit any of the steps in the formation of both UDPNAG and UDPNAM-pentapeptide. However, inhibition of the synthesis of the latter is likely to be potentially more rewarding since its formation requires a larger number of steps, which gives a wider scope for intervention. Drugs in clinical use that act by inhibiting different processes in this stage of cell wall synthesis are cycloserine and fosfomycin (Fig.).

Cycloserine and fosfomucin

 

An outline of the biosynthesis of the precursors of the peptidoglycan chain of the cell wall of Staphylococcus aureus

Cycloserine is a broad spectrum antibiotic produced by Streptomyces orchidaceus. The drug

is used mainly as a second-line antitubercular agent. It enters the bacteria by active transport systems, which results in a high concentration in the bacterial cell, a primary requirement for activity. D-Cycloserine inhibits both alanine racemase and D-alanyl-D-alanine synthetase, which blocks the conversion of the tripeptide to the pentapeptide at two places (A in Fig.). The affinity of the enzymes in Staphylococcus aureus for the drug has been found to be 100 times higher than its affinity for its natural substrate D-alanine. This affinity is believed to depend on the isoxazole ring, whose shape corresponds to one of the conformations of D-alanine. It is believed that the rigid structure of the isoxazole ring gives the drug a better chance of binding to the active sites of the enzymes than the more flexible structure of D-alanine.

 

An outline of the formation of the peptidoglycan chains of Staphylococcus aureus from UDPNAM-pentapeptide and UDPNAG. UMP is uridine monophosphate

 

 

Fosfomycin, produced by a number of Streptomyces species, is active against both Gram-positive and Gram-negative bacteria. However, it is used mainly to treat Gram-positive infections. The drug acts by inhibiting the enol-pyruvate transferase (B in Fig.) that catalyses the incorporation of phosphoenolpyruvic acid (PEP) into the UDPNAG molecule. However, the drug does not inhibit other enol-pyruvate transferases used to incorporate PEP in a number of other biosynthetic reactions. Consequently, it appears that the activity of the drug is due to it forming an inactive product with the enzyme. It has been suggested that this product is formed by the acid-catalysed nucleophilic substitution of the oxiran ring by the sulphydryl groups of the cysteine residues in the active site of the enzyme.

 

Drugs that inhibit the synthesis of the peptidoglycan chain The sequence of reactions starting from UDPNAM-pentapeptide and UDPNAG to form the peptidoglycan chain (Fig.) is not completely known although the main stages have been identified. However, it is known that the reactions are catalysed by membrane-bound enzymes. A number of antibiotics, such as bacitracin, are believed to inhibit some of the stages of the biosynthesis of the peptidoglycan chains.

 

Bacitracin is a mixture of similar peptides produced from Bacillus subtilis. The main component of this mixture is bacitracin A, which is active against Gram-positive bacteria. However, its high degree of neuro- and nephrotoxicity means that the drug is seldom used, and then somewhat cautiously. Its main site of action appears to be inhibition of the dephosphorylation of membrane-bound phospholipid carrier bactoprene (step M in Fig.). Its action is enhanced by the presence of zinc ions.

 

 

Drugs that inhibit the cross-linking of the peptidoglycan chains The final step in the formation of the cell wall is the completion of the cross-links. This converts the water-soluble and therefore mobile peptidoglycans into the insoluble stationary cell wall. Investigations using Staphylcoccus aureus indicated that the cross-linking is brought about by a multistep displacement of the terminal alanine of the peptide attached to one peptidoglycan chain and its replacement by the terminal glycine of the peptide attached to a second peptidoglycan chain (Fig.). This reaction is catalysed by transpeptidases.

 

An schematic outline of the formation of the peptide cross-links in the formation of the cell wall of Staphylcoccus aureus

 

The b-lactam group of antibiotics inhibit cell wall synthesis by inhibiting the transpeptidases responsible for the cross-linking between the peptidoglycan chains. This group of antibiotics, named after the b-lactam ring that they all have in common, includes the widely used penicillins and the cephalosporins (Fig.). Both of these groups of beta-lactam antibiotics are more effective against Gram-positive bacteria than Gram-negative bacteria. However, some cephalosporins, such as ceftazidime, which is administered intravenously, are very effective against Gram-negative bacteria.

 

Examples of the range of penicillins and cephalosporins. The R residues of ampicillin, amoxicillin and ceftazidime have D configurations

 

The b-lactam antibiotics have to reach the plasma membrane of the bacteria before they can act. As the outer surfaces of Gram-positive bacteria are covered by a thin layer of teichoic acids. they offer less resistance to drug penetration than Gram-negative bacteria, where the drug has to penetrate both the outer membrane and the periplasmic space (Fig.) before it can interfere with cell wall synthesis. In Gram-negative bacteria the drug diffuses through the outer membrane via porin channels formed by integral trimeric proteins. A large number of these channels, with diameters of about 1.2 nm, are found in each bacterial cell wall. However, not all porin channels are able to transport b-lactam drugs. Some bacterial genera such as Pseudomonas are resistant to b-lactam antibiotics because their porin channels will not allow the transport of these drugs.

 

Once through the outer membrane the drug diffuses across the periplasmic space, which contains b-lactamases that can inactivate the drug. Gram-positive bacteria also produce these enzymes, which they release into the extracellular fluid. Finally, the drug penetrates to the outer surface of the plasma membrane where it binds to, and blocks the action of, the transpeptidases and other proteins involved in cell wall synthesis. The precise nature of the blocking mechanism has not yet been fully elucidated but appears to involve the b-lactam ring system. This ring system is very reactive and is easily decomposed by acid and base catalysed hydrolysis (Fig.), the rate of which depends on the structure of the penicillin.

 

 

Some of the decomposition routes of the b-lactam ring of benzylpenicillin

The final stage in the formation of the cross-links between the peptidoglycan chains in bacteria is catalysed by a glycopeptide transpeptidase. It is believed that the hydroxyl group of a serine residue in this enzyme displaces the last alanine residue from the tetrapeptide chain. The displaced alanine diffuses away from the reaction site, allowing attack by the amino group of the terminal glycine of the pentaglycine chain on the alanine bound to the enzyme to complete the peptide linkage and regenerate the enzyme (Fig.). However, it is thought that since the geometry of the penicillins resembles that of the alanyl-alanyl unit the bacteria mistakes the drug for its normal substrate. The b-lactam ring reacts with the enzyme to form a covalently bound acyl derivative. The 1,3-thiazolidine ring of this derivative is believed to prevent a pentaglycine unit from attacking the enzyme–acyl linkage and regenerating the active site of the enzyme.

 

A schematic outline of the chemistry proposed for the action of penicillins

 

Penicillins are unstable under acid conditions, the rate of decomposition depending on which penicillin is being considered. For example, piperacillin (Fig.) is so acid labile that it has to be administered by intravenous infusion. This means that the acid conditions of the stomach could reduce the amount of the drug reaching the general circulatory system and hence its effectiveness. The reactivity of penicillins under acid conditions can be attributed to the presence of a reactive four-membered lactam ring, the carbonyl group of which is readily attacked by nucleophiles under acid conditions.

This reactivity is believed to be enhanced by the presence of the acyl side chain in a so-called neighbouring group effect. This group is believed to enhance the electronegative nature of the oxygen of the carbonyl group of the lactam, which makes the carbon of the lactam group more susceptible to nucleophilic attack (Fig.).

A possible mechanism for the enhancement of the reactivity of the carbonyl group of the lactam by a neighbouring group

Consequently, some acid-resistant penecillins have been produced by introducing an electron withdrawing group on the alpha carbon of the side chain. This reduces neighbouring group participation and, as a result, the reactivity on the lactam’s carbonyl group (Fig.). This approach led to the development of penicillin V (phenoxymethyl-penicillin), amoxacillin and ampicillin.

Cephalosporins usually exhibit a greater resistance to acid hydrolysis than penicillins. However, the first generation of cephalosporins were not as potent as the penicillins but were

active against a wider range of bacteria. However, their absorption from the GI tract is often

poor and so they have to be given by injection. Consequently, cephalosporin C, first isolated by workers in Oxford University in the late 1940s, was used as the lead to develop more active

analogues (Fig.). A large number of different cephalosporins are now in clinical use. The relationship between the structures of b-lactams and their activity has been the subject of much discussion. Originally it was believed that the amide-linked side chain, the carboxylic acid at position 2 and the fused thio-ring systems were all essential for the pharmacological activity of b-lactam antibiotics. However, the discovery of b-lactams such as thienamycin, aztreonam and nocardin A (Fig.), whose structures do not contain all these functional groups, suggests that the b-lactam ring is the only essential requirement for activity.

 

(a) It is believed that the electron withdrawing effect of the R group reduces the ability of the electrons of the carbonyl group of the amide link to influence those of the carbonyl group of the lactam. (b) Examples of penicillins in clinical use with an electron withdrawing R group

The increasing number of bacteria resistant to b-lactam antibiotics is becoming a major problem. Resistance to penicillins and cephalosporins by some bacteria is mainly due to inactivation of the drug by hydrolysis of the lactam ring catalysed by the b-lactamases produced by that bacterium. However, in general, penicillins tend to be more susceptible than cephalosporins to hydrolysis catalysed by b-lactamases.

 

Both Gram-positive and Gram-negative bacteria produce b-lactamases. In the former case the enzyme is liberated into the medium surrounding the bacteria. This results in inactivation of the penicillin, cephalosporin and other b-lactam drugs before the drug reaches the bacteria. However, with Gram-negative bacteria, the hydrolysis takes place within the periplasmic space. In addition some Gram-negative bacteria produce Acylases which can cleave the side chains of penicillins. Bacteria that have developed a resistance to b-lactam antibiotics are often treated using a dosage form incorporating a b-lactamase inhibitor such as clavulanic acid, sulbactam or tazobactam (Fig.) and Table.

 

Examples of b-lactam antibiotics that do not contain a thiol-ring system

 

Examples of b-lactamase inhibitors used in penicillin dosage forms

 

 

Examples of dosage forms containing b-lactamase inhibitors

An alternative approach to the problem of bacterial resistance was the development of b-lactamase-resistant drugs. The strategy adopted by Beecham Research Laboratories for penicillins in 1961 was to use bulky substituents close to the labile lactam ring in order to use

steric hindrance to prevent the drug binding to the enzyme’s active site and undergoing lactam hydrolysis. This approach resulted in the discovery of methicillin. However, methicillin had a reduced potency compared with other penicillins and was acid labile to the extent that it could only be administered by injection. Further work by Beecham using an isoxazole ring in the side chain resulted in the discovery of flucloxacillin (Fig.), oxacillin, cloxacillin and dicloxacillin. These drugs are used against Gram positive bacteria. They are inactive against Gram negative bacteria. However, oxacillin is also acid resistant.

 

The introduction of a bulky syn a-oximino group (-C-N-O-) side chain in so-called ‘second-generation cephalosporins’ improved their stability towards b-lactamases and esterases by probably sterically hindering the hydrolysis of the lactam. For example, cefuroxime (Fig.) is active against a wide range of Gram-positive and Gram-negative bacteria. Further development has resulted in the discovery of so-called ‘third- and fourth-generation’ cephalosporins that also exhibit enhanced resistance to b-lactamases. This increased resistance is believed to be partly due to the presence of a syn a-oximino side chain. In the third-generation cephalosporins, such as ceftazidine (Fig.), the presence of an aminothiazole group is thought to increase the ease of transfer of the drug through the outer membrane of Gram-negative bacteria, which would account for their good Gram-negative activity and variable Gram-positive activity. The fourth-generation cephalosporins, such as cefepime and cefpirome, are zwitterions. They have a high potency against Gram-positive, Gram-negative and Pseudomonas aeruginosa.

 

Polypeptide antibiotics A large number of polypeptide antibiotics have been discovered. They are active against a wide variety of microorganisms and operate by a range of mechanisms. Vancomycin and teicoplanin (teichomycin A2) act by inhibiting the cross-linking of the peptidoglycan chains in bacterial cell walls. Other polypeptide antibiotics such as gramicidin and viomycin, which is used to treat TB, have different mechanisms of action. Vancomycin (Fig.a) is a glycopeptide antibiotic that was isolated from Streptomyces orientalis in 1955 and inhibits the formation of the peptide links between the peptidoglycan chains. In spite of the extensive use of the drug, very little bacterial resistance to vancomycin has developed. It is mainly used for Gram-positive infections but is irritating on intravenous injection. Oral administration does not give useful blood levels. However, the drug is used orally to treat pseudomembranous enterocolitis caused by high concentrations of Clostridium difficile in the intestine.

 

The structure of vancomycin (Fig.a) is based on a tricyclic ring system of aliphatic and aromatic amino acids with a disaccharide side chain. This structure is rigid with a peptide-lined pocket that has a strong affinity for D-ala-D-ala residues. Vancomycin inhibits cell wall synthesis by binding to the D-ala-D-ala end group of the pentapeptide chain of the peptidoglycan cell wall precursor. NMR spectroscopy and molecular modelling suggest that the D-ala-D-ala residue is multiple hydrogen-bonded to the vancomycin in this pocket. This inhibits the formation of the peptide cross-links between the polyglycan chains (Fig.), which results in a loss of bacterial cell wall integrity.

 

Those bacteria that exhibit resistance to the drug appear to have replaced the D-ala-D-ala unit of the pentapeptide residue by a D-ala-D-lactate unit. This structural change is believed to reduce the number of hydrogen bonds between the drug and the D-ala-D-lactate of the peptidoglycan precursor by one. However, this small change is sufficient to prevent the drug operating efficiently. Teicoplanin is a complex of five compounds (A2-1, A2-2 to A2-5) with similar structures (Fig. b) that is active against a range of Gram-positive and Gram-negative bacteria.

It was isolated in 1976 from Actinoplanes teichomycetius. The components of the mixture were separated in 1983 and their structures elucidated in 1984. Teicoplanin has good water solubility but significantly better lipid solubility than vancomycin. It is generally less toxic than vancomycin and its long half-life means that it need only be administered once a day. Its mechanism of action is believed to be identical to that of vancomycin.

 

Surfactants used as antibacterial agents

Surface active agents disrupt cell membranes because they dissolve in both the aqueous extracellular fluid and the lipid membrane. This lowers the surface tension of the membrane, which allows water to flow into the cell and ultimately results in lysis and bactericidal action. In all cases a balance between the hydrophilic and lipophilic sections of the molecule is essential for action. Both cationic and non-ionic surfactants are used (Fig). In addition, detergent surfactants, such as sodium dodecyl sulphate, are also used to remove proteins from cell membranes.

 

Chloramphenicol

Chloramphenicol was first isolated by Ehrlich et al. in 1947 from the microorganism Streptomyces venezuelae, which was found in a soil sample from Venezuela. It is a broad spectrum antibiotic whose structure contains two asymmetric centres. However, only the D-(-)-threo form is active. Its use can cause serious side effects and so it is recommended that chloramphenicol is only used for specific infections. It is often administered as its palmitate in order to mask its bitter taste. The free drug is liberated from this ester by esterase-catalysed hydrolysis in the duodenum. Chloramphenicol has a poor water solubility (2.5 g dm-3) and so it is sometimes administered in the form of its more soluble sodium hemisuccinate salt, which acts as a prodrug. Chloramphenicol is believed to act by inhibiting the elongation stage in protein synthesis in prokaryotic cells. It binds reversibly to the 50S ribosome subunit and is thought to prevent the binding of the aminoacyl–tRNA complex to the ribosome. However, its precise mode of action is not understood.

 

Investigation of the activity of analogues of chloramphenicol showed that activity requires a para-electron withdrawing group. However, substituting the nitro group with other electron withdrawing groups gave compounds with a reduced activity. Furthermore, modification of the side chain, with the exception of the difluoro derivative, gave compounds that had a lower activity than chloramphenicol (Table). These observations suggest that D-(-)-threo-chloramphenicol has the optimum structure of those tested for activity.

The activity against E. coli of some analogues of chloramphenicol relative to chloramphenicol

The synthesis of chloramphenicol was first reported by Controulis J et al. in 1949 (Fig.). Numerous synthetic routes have since been devised for the synthesis of chloramphenicol, the commercial routes usually starting with 4-nitroacetophenone. Chloramphenicol is now manufactured by both totally synthetic and microbiological routes.

 

Antiviral drugs

It has been found that viruses utilise a number of virus-specific enzymes during replication. These enzymes and the processes they control are significantly different from those of the host cell to make them a useful target for medicinal chemists. Consequently, antiviral drugs normally act by inhibiting viral nucleic acid synthesis, inhibiting attachment to and penetration of the host cell or inhibiting viral protein synthesis.

 

An outline of the mechanism for DNA chain termination used by some antiviral drugs. The growing chain terminates because R cannot react with the next triphosphate nucleotide (for DNA replication, see section 10.5). Key: P = phosphate residue and R = hydrogen and various other groups (Table)

Nucleic acid synthesis inhibitors

Nucleic acid synthesis inhibitors usually act by inhibiting the polymerases or reverse transcriptases required for nucleic acid chain formation. However, because they are usually analogues of the purine and pyrimidine bases found in the viral nucleic acids, they are often incorporated into the growing nucleic acid chain. In this case their general mode of action frequently involves conversion to the corresponding 5-triphosphate by the host cell’s cellular kinases. This conversion may also involve specific viral enzymes in the initial monophosphorylation step. These triphosphate drug derivatives are incorporated into the nucleic acid chain where they terminate its formation. Termination occurs because the drug residues do not have the 30-hydroxy group necessary for the phosphate ester formation required for further growth of the nucleic acid chain. This effectively inhibits the polymerases and transcriptases that catalyse the growth of the nucleic acid (Fig.).

 

It is not possible to list all the known antiviral agents in this text so only a representative selection are discussed.

 

Aciclovir

Aciclovir was the first effective antiviral drug. It is effective against a number of herpes viruses, notably simplex, varicella-zoster (shingles), varicella (chickenpox) and Epstein–Barr virus (glandular fever). It may be administered orally and by intravenous injection as well as topically. Orally administered doses have a low bioavailability.

The action of aciclovir is more effective in virus-infected host cells because the viral thymidine kinase is a more efficient catalyst for the monophosphorylation of aciclovir than the thymidine kinases of the host cell. This leads to an increase in the concentration of the aciclovir triphosphate, which has 100-fold greater affinity for viral DNA polymerase than human DNA polymerase. As a result, it preferentially competitively inhibits viral DNA polymerase and so prevents the virus from replicating. However, resistance has been reported due to changes in the viral mRNA responsible for the production of the viral thymidine kinase. Aciclovir also acts by terminating chain formation. The aciclovir–DNA complex formed by the drug also irreversibly inhibits DNA polymerase.

 

Vidarabine

Vidarabine is active against herpes simplex and herpes varicella-zoster. However, the drug does give rise to nausea, vomiting, tremors, dizziness and seizures. In addition it has been reported to be mutagenic, teratogenic and carcinogenic in animal studies. Vidarabine is administered by intravenous infusion and topical application. It has a half-life of about one hour, the drug being rapidly deaminated to arabinofuranosyl hypoxanthine (ara-HX) by adenosine deaminase. This enzyme is found in the serum and red blood cells. Ara-HX, which also exhibits a weak antiviral action, has a half-life of about 3.5 hours.

 

Ribavirin

Ribavirin is effectively a guanosine analogue. It is active against a wide varietyof DNA and RNAviruses but the mechanism by which it acts is not understood. It is mainly used in aerosol form to treat influenza and other respiratory viral infections. Intravenous administration in the first 6 days of onset has been effective in reducing deaths from Lassa fever to 9 per cent. Ribavirin has also been shown to delay the onset of full-blown AIDS in patients with early symptoms of HIV infection. However, administration of the drug has been reported to give rise to nausea, vomiting, diarrhoea, deterioration of respiratory function, anaemia, headaches and abdominal pain. The mechanism by which it acts may differ from one virus to another.

Zidovudine (AZT)

Zidovudine was originally synthesised in 1964 as an analogue of thymine by J. Horwitz as a potential antileukaemia drug. It was found to be unsuitable for use in this role and for 20 years was ignored, even though in 1974W. Osterag et al. reported that it was active against Friend leukaemia virus, a retrovirus. However, the identification in 1983 of the retrovirus HIVas the source of AIDS resulted in the virologist M. St Clair setting up a screening programme for drugs that could attack HIV. Fourteen compounds were selected and screened against Friend leukaemia virus and a second retrovirus called Harvey sarcoma virus. This screen led to the discovery of zidovudine (AZT), which was rapidly developed into clinical use on selected patients in 1986.

AZT is converted by the action of cellular thymidine kinase to the 50-triphosphate. This inhibits the enzyme reverse transcriptase in the retrovirus, which effectively prevents it from forming the viral DNA necessary for viral replication. The incorporation of AZT into the nucleic acid chain also results in chain termination because the presence of the 30-azide group prevents the reaction of the chain with the 50-triphosphate of the next nucleotide waiting to join the chain (Fig.). AZT is also active against mammalian DNA polymerase and although its affinity for this enzyme is about 100-fold less this action is thought to be the cause of some of its unwanted side effects.

 

Zidovudine is active against the retroviruses that cause AIDS (HIV virus) and certain types of leukaemia. It also inhibits cellular a-DNA polymerase but only at concentrations in excess of 100-fold greater than those needed to treat the viral infection. The drug may be administered orally or by intravenous infusion. The bioavailability from oral administration is good, the drug being distributed into most body fluids and tissues. However, when used to treat AIDS it has given rise to gastrointestinal disorders, skin rashes, insomnia, anaemia, fever, headaches, depression and other unwanted effects. Resistance increases with time. This is known to be due to the virus developing mutations’ which result in changes in the amino acid sequences in the reverse transcriptase.

Didanosine

Didanosine is used to treat some AZT-resistant strains of HIV. It is also used in combination with AZT to treat HIV. Didanosine is administered orally in dosage forms that contain antacid buffers to prevent conversion by the stomach acids to hypoxanthine. However, in spite of the use of buffers the bioavailability from oral administration is low. The drug can cause nausea, abdominal pain and peripheral neuropathy, amongst other symptoms. Drug resistance occurs after prolonged use. Didanosine is converted by viral and cellular kinases to the monophosphate and then to the triphosphate. In this form it inhibits reverse transcriptase and in addition its incorporation into the DNA chain terminates the chain because the drug has no 30-hydroxy group (Fig.).

 

Host cell penetration inhibitors

The principal drugs that act in this manner are amantadine and rimantadine (Fig.). Both amantadine and rimantadine are also used to treat Parkinson’s disease. However, their mode of action in this disease is different from their action as antiviral agents.

 

Examples of host cell penetration inhibitors

 

 

Amantadine hydrochloride

Amantadine hydrochloride is effective against influenza A virus but is not effective against the influenza B virus. When used as a prophylactic, it is believed to give up to 80 per cent protection against influenza A virus infections. The drug acts by blocking an ion channel in the virus membrane formed by the viral proteinM2. This is believed to inhibit the disassembly of the core of the virion and its penetration of the host. Amantadine hydrochloride has a good bioavailability on oral administration, being readily absorbed and distributed to most body fluids and tissues. Its elimination time is 12–18 hours. However, its use can result in depression, dizziness, insomnia and gastrointestinal disturbances, amongst other unwanted side effects.

 

 

Rimantadine hydrochloride

Rimantadine hydrochloride is an analogue of amantadine hydrochloride. It is more effective against influenza A virus than amantadine. Its mode of action is probably similar to that of amantadine. The drug is readily absorbed when administered orally but undergoes extensive first-pass metabolism. However, in spite of this, its elimination half-life is double that of amantadine. Furthermore, CNS side effects are significantly reduced.

 

Inhibitors of viral protein synthesis

The principal compounds that act as inhibitors of protein synthesis are the interferons. These compounds are members of a naturally occurring family of glycoprotein hormones (RMM 20 000–160 000), which are produced by nearly all types of eukaryotic cell. Three general classes of interferons are known to occur naturally in mammals, namely:

the a-interferons produced by leucocytes, b-interferons produced by fibroblasts and g-interferons produced by T lymphocytes. At least twenty a-, two b- and two g-interferons have been identified. Interferons form part of the human immune system. It is believed that the presence of virions, bacteria and other antigens in the body switches on the mRNA that controls the production and release of interferon. This release stimulates other cells to produce and release more interferon. Interferons are thought to act by initiating the production in the cell of proteins that protect the cells from viral attack. The main action of these proteins takes the form of inhibiting the synthesis of viral mRNA and viral protein synthesis. a-Interferons also enhance the activity of killer T cells associated with the immune system.

A number of a-interferons have been manufactured (see Table 10.10 on page 394) and proven to be reasonably effective against a number of viruses and cancers. Interferons are usually given by intravenous, intramuscular or subcutaneous injection. However, their administration can cause adverse effects, such as headaches, fevers and bone marrow depression, that are dose related.

The formation and release of interferon by viral and other pathological stimulation has resulted in a search for chemical inducers of endogenous interferon. Administration of a wide range of compounds has resulted in the induction of interferon production. However, no clinically useful compounds have been found for humans’ although tilorone is effective in inducing interferon in mice.

Antimalarial compounds

The causative agents of malaria are plasmodia, unicellular organisms (Order Hemosporidia, Class Protozoa). The infective form, the sporozoite, is inoculated into skin capillaries when infected female Anopheles mosquitoes (A) suck blood from humans. The sporozoites invade liver parenchymal cells, where they develop into primary tissue schizonts. These give rise to numerous merozoites that enter the blood. The preerythrocytic stage is asymptomatic. In blood, the parasite enters erythrocytes (erythrocytic stage), where it again multiplies by schizogony, resulting in the formation of more merozoites. Rupture of the infected erythrocytes releases the merozoites and pyrogens. A fever attack ensues and more erythrocytes are infected. The generation period for the next crop of merozoites determines the interval between fever attacks. With Plasmodium vivax and P. ovale, there can be a parallel multiplication in the liver (paraerythrocytic stage).

 

 

Moreover, some sporozoites may become dormant in the liver as “hypnozoites” before entering schizogony.  Different antimalarials selectively kill the parasite’s different developmental forms. The mechanism of action is known for some agents: Chloroquine and quinine accumulate within the acidic vacuoles of blood schizonts and inhibit polymerization of heme released from digested hemoglobin, free heme being toxic for the schizonts. Pyrimethamine inhibits protozoal dihydrofolate reductase, as does chlorguanide (proguanil) via its active metabolite cycloguanil. The sulfonamide sulfadoxine inhibits synthesis of dihydrofolic acid. Dihydrofolate reductase is also blocked by cycloguanil, the active form of proguanil. Atoquavone suppresses synthesis of pyrimidine bases, probably by interfering with mitochondrial electron transport. Artemesinin derivatives (artemether, artesunate) originate from the East Asian plant Qinghaosu (Artemisia sp.) Its antischizontal effect appears to involve a reaction between heme iron and the epoxide group of these compounds. Antimalarial drug choice takes tolerability and plasmodial resistance into account.

 

Tolerability

The oldest antimalarial, quinine, has the smallest therapeutic margin. All newer agents are rather well tolerated. Plasmodium falciparum, responsible for the most dangerous form of malaria, is particularly prone to develop drug resistance. The prevalence of resistant strains rises with increasing frequency of drug use. Resistance has been reported for chloroquine and also the combination pyrimethamine/sulfadoxine.

 

Drug choice for antimalarial chemoprophylaxis

In areas with a risk of malaria, continuous intake of antimalarials affords the best protection against the disease, though not against infection. Primaquine would be effective against primary tissue schizonts of all plasmodial species; however, it is not used for long-term prophylaxis because of unsatisfactory tolerability and the risk of plasmodial resistance. Instead, prophylactic regimens employ agents against blood schizonts. Depending on the presence of resistant strains, use can be made of chloroquine, and/or proguanil, mefloquine, the tetracycline doxycycline, as well as the combination of atoquavone and proguanil. These drugs do not prevent the (symptom-free) hepatic infection but only the disease-causing infection of erythrocytes (“suppression therapy”). On a person’s return from an endemic malaria region, a two-week course of primaquine is adequate for eradication of the late hepatic stages (P. vivax and P. ovale). Protection frommosquito bites (with nets, skin-covering clothes, etc.) is a very important prophylactic measure.

 

Therapy

Antimalarial therapy employs the same agents, in addition to the combinations of artemether plus lumefantrine or pyrimethamine plus sulfadoxine.

 

Enzymes and drug resistance

Drug resistance occurs when a drug no longer has the desired clinical effect. This may be due to either a natural inbuilt resistance in some individuals and organisms or may arise naturally in the course of treatment. The former is probably due to differences in the genetic code of individuals within a species whilst the latter arises because of natural selection. In natural selection the drug kills the susceptible strains of an organism but does not affect other strains of the same organism. Consequently, these immune strains multiply and become the common strain of the organism, which subsequently results in ineffective drug treatment. Resistance occurs on an individual basis and so is not usually detected until a wide sample of the population has been treated with or indirectly exposed to the drug. Its detection necessitates the discovery of new drugs to treat the condition. Its emergence is probably due to the widespread and poorly controlled use of a drug. For example, the generous use of antibiotics in farming is strongly suspected to be the reason for an increase in antibiotic-resistant strains of bacteria in humans. The response of medicinal chemists to resistance is either to devise new drugs or to modify existing drugs. This approach suffers from the high probability of being unsuccessful as well as time consuming and expensive. In the light of human experience it would be better if, in future, we reduced the possibility of resistance by using the effective existing drugs more intelligently. Drug resistance can be linked to a change in either the permeability of the membranes of the organism or an enzyme system(s) of the organism. Enzymes may be involved in drug resistance in a number of ways but in many cases resistance may be due to several different processes occurring at approximately the same time.

 

Changes in enzyme concentration

A significant increase or decrease from the normal concentration of an enzyme can result in resistance to a drug. The overproduction of an enzyme can have two effects:

1. The target process catalysed by the enzyme will not be inhibited because excess enzyme is produced. For example, the resistance of malarial parasites is believed to be caused by overproduction of dihydrofolate reductase due to the drug stimulating the parasite’s RNA.

 

2. The increased production of enzymes that inactivate the drug, for example beta-lactamases inactivate most penicillins and cephalosporins by hydrolysing their b-lactam rings. A number of enzymes deactivate inhibitors by incorporating (conjugation) phosphate by phosphorylation of hydroxyl groups, adenine by adenylation of hydroxyl groups or acetyl by acetylation of amino groups in the inhibitor’s structure (Fig.). ATP is believed to be the usual provider of phosphate and adenylic acid, whilst acetyl coenzyme A is thought to be the normal source of acetyl groups. For example, resistance to the antibiotic kanamycin A can occur by all three routes (Fig.) although kanamycin A-resistant bacteria do not normally use all three routes.

 

The inhibition of kanamycin A by enzymatic inactivation. The arrows indicate the structure and position of the result of an enzyme reaction

 

 

Many aminoglycoside antibiotics are susceptible to this type of enzyme inhibition. However, amikacin (AK) where the C1-NH2 of kanamycin A has been acylated by S-a-hydroxy-gamma-aminobutanoic acid is not susceptible to 3-phosphorylation and 2-adenylation. The underproduction of an enzyme could result in insufficient enzyme being present to produce the active form of a drug from a prodrug. For example the resistance to the antileukaemia drug 6-mercaptopurine is caused by a reduced production of hypoxanthine–guanine phosphoribosyltransferase, the enzyme required to convert the prodrug to its active ribosyl 5-monophosphate derivative.

These changes in the production of the enzyme are believed to be due to genetic changes in the organism.

 

An increase in the production of the substrate

Increased production of the substrate can prevent competitive reversible inhibitors from binding to the active site in sufficient quantities to be effective. The high concentration of the substrate moves the position of equilibrium to favour the formation of the E–S complex.

For example, the inhibition of dihydropteroate synthetase by sulphonamides results in a build-up of p-aminobenzoic acid. This increase in substrate concentration prevents sulphonamides from inhibiting dihydropteroate synthetase, which is a key enzyme in the production of the RNA necessary for bacterial reproduction. Similarly, a build-up of angiotensin I will overcome the effect of ACE inhibitors. It is thought to be the reason for the concentration of plasma angiotensin II returning to normal in some cases where there has been a chronic administration of ACE inhibitors.

 

Changes in the structure of the enzyme

Changes in the structure of the target enzyme result in a structure that is not significantly inhibited by the drug. However, the modified enzyme is still able to produce the normal product of the reaction, which allows the unwanted metabolic pathway to continue to function. For example, resistance to the antibiotic trimethoprim is believed to be due to a plasmid-directed change in the structure of dihydrofolate reductase in the bacteria.

 

Similarly, the resistance of the E. coli strains to sulphonamides has been shown to be due to their containing a sulphonamide-resistant dihydropteroate synthase.

 

The use of an alternative metabolic pathway

The blocking of a metabolic pathway by a drug can result in the opening of a new pathway controlled by a different enzyme that is not inhibited by the same drug.

 

 

 

Agonist, partial agonist, inverse agonist, biased agonist and antagonist

Agonist: A drug that mimics the endogenous receptor ligand to activate the receptor to produce a biological response is called as an agonist. Several agonists are able to produce the target maximum response without completely occupying all the receptors.

Partial agonist: A drug that binds and activates a receptor but does not elicit a full response is known as a partial agonist. A partial agonist can block the effect of a full agonist. In the presence of high concentrations of a partial agonist, the action of a full agonist can be reduced to the maximum response elicited by the partial agonist. However, the intrinsic activity would be greater than zero but less than 1 that of a full agonist.

Inverse agonist: An inverse agonist is a molecule or agent that binds to the same receptor site as an agonist and is considered to be a full agonist. However, it exerts the opposite pharmacological response to that of a normal agonist, i.e. demonstrates negative efficacy. Constitutive activity refers to the ability of a receptor in producing its biological response in the absence of a bound ligand. The constitutive activity of a receptor may be blocked by an inverse agonist.

Biased agonist: G protein-coupled receptors (GPCRs) are capable of signalling with different efficacies to their multiple downstream pathways, a phenomenon referred to as biased agonism. Biased agonism is one of the fastest growing areas in GPCR pharmacology. Biased agonism has been primarily reported as a phenomenon of synthetic ligands and the biological importance of such signalling is unclear.

Antagonist: A drug that binds to a receptor but does not elicit a response is referred to as an antagonist. Importantly, the antagonist must block the action of the agonist at the receptor site. Antagonist can shift the concentration–response curve of an agonist to the right by reducing its fractional occupancy. High concentrations of the antagonist may block the actions of the agonist completely. However, antagonists have no intrinsic activity and therefore they do not produce any effects. There are two main types of antagonists. Competitive antagonists compete with the agonist for same receptor binding site, but the binding is reversible. It shifts the concentration-response curve of the agonist to the right without any reduction in maximal response. Non-competitive antagonists bind irreversibly to a receptor site and thereby reduce the ability of an agonist to bind and produce a response. The non-competitive antagonism is a slow process which resulting in a prolonged antagonistic effect.

Drug affinity and efficacy

Affinity can be defined as the extent or fraction to which a drug binds to receptors at any given drug concentration or the firmness with which the drug binds to the receptor. The mathematical model of affinity of a drug for the receptor was first described by Irving Langmuir Kenakin (2004). Affinity is one of the factors that determine potency. Affinity is inversely proportional to the potency of a drug (1Kd), where Kd is the dissociation constant. The strength of the binding (interaction) of a ligand and its receptor can be described by affinity. The higher the Kd value, the weaker the binding and the lower the affinity. The opposite occurs when a drug has a low Kd. Potency is a measure of necessary amount of the drug to produce an effect of a given magnitude. In general, potency is denoted as the median effective concentration/dose as EC50/ED50/Kd.

Efficacy (intrinsic activity) is the ability of a drug to illicit a pharmacological response (physiological) when interaction occurs with a receptor (relationship between response and occupancy of receptor). Efficacy depends on the efficiency of the receptor activation to cellular responses and the formation of number of drug-receptor complexes.

•Full agonists: efficacy = 1.

•Partial agonists: efficacy > 0 and < 1.

•Competitive antagonists: efficacy = 0.

 

Clark’s occupancy theory

Clark in the 1920s visualised the drug–receptor interaction as being a bimolecular dynamic

equilibrium with the drug molecules continuously binding to and leaving the receptor, that is:

Clark stated that the intensity of the response at any time was proportional to the number of

receptors occupied by the drug: the greater the number occupied, the greater the

pharmacological effect, that is:

According to Clark a maximum response would be obtained when all the receptors were

occupied, that is:

 

where RT is the total number of receptors. It follows from equations (8.8) and (8.9) that for

a given dose of a drug the fraction of the maximum response is given by:

The dissociation of the drug–receptor complex may be represented as:

and applying the law of mass action:

where KD is the dissociation constant for the drug–receptor complex.

But the total receptor concentration is:

Substituting equation (8.13) in equation (8.12):

 

Equation (8.19) shows that the relationship between E and molar drug concentration [D] is in the form of a rectangular hyperbola, whilst that between E and log [D] is sigmoidal. These theoretical relationships derived using Clarks’ theory are often in good agreement with the experimental results (Fig.) obtained in a number of investigations. Furthermore, substituting the value of E/Emax = 1/2 in equation (8.19) gives the relationship:

where EC50 is the molar concentration of the drug that produces half the maximum biological response observed when a ligand binds to a receptor. However, in practice this theoretical relationship does appear to be the exception rather than the rule. The value of the dissociation constant KD is a measure of the affinity of the drug for the receptor. Drugs with small KD values have a large affinity for the receptor whilst those with high values have a low affinity. As a result, KD values are used to compare the activities of a series of analogues during drug development. The value of KD may be determined experimentally from tissue binding experiments using a radioactive form of the drug. The data obtained from this type of experimental work may be analysed using a Scatchard plot of the ratio of bound to free ligand against bound drug (Fig.). This gives a straight line with a slope of -1/KD provided that the drug binds to only one type of receptor. In industry the data are now analysed by the use of a computerised method of least squares.

 

The correlation of experimental results and those predicted using Clark’s theory for the stimulated contraction of guinea-pig ileum by acetylcholine

 

Although Clark’s occupancy theory is still a cornerstone of pharmacodynamics, a number of its assumptions have now been shown to be incorrect. It is now known that:

·       the formation of many drug–receptor complexes is not reversible;

·       the receptor sites are not always independent;

·       the formation of the complex may not be bimolecular: for example, two acetylcholine

molecules bind to nACh receptors of ion channels;

·       a maximum response may be obtained before all the receptors are occupied;

·       the response is not linearly related to the proportion of receptors occupied, especially

in the case of partial agonists.

 

In the 1950s. Ariens and Stephenson separately modified Clark’s theory to account for the existence of agonists, partial agonists and antagonists. They based their modifications on a proposal by Langley in 1905, which visualised the action of a receptor as taking place in two stages. The first stage was the binding of the ligand to the receptor, which was controlled by the ligand’s affinity for the receptor. The second stage was the initiation of the biological response. Ariens said that this second step was governed by the ability of the ligand–receptor complex to initiate a response. Ariens called this ability the intrinsic activity (a), whilst Stephenson referred to it as the efficacy (e) of the ligand–receptor complex. Intrinsic activity may be defined as:

Using the concept of intrinsic activity Clark’s equation (equation 8.19) becomes:

When a = 1 for ligands with identical affinities for a receptor, equation (8.22) reverts to the original form of Clark’s equation (equation 8.19). This means that a normal response curve is obtained and the drug acts as a full agonist. However, when a = 0 the percentage response is zero and the drug is a full antagonist. Intermediate values between 1 and 0 for a indicate a partial agonist (Fig.).

In 1950s, Stephenson discovered that a maximum response was obtained when only a proportion of the available receptors were occupied. This discovery was in direct conflict with Clark’s occupancy theory and led Stephenson to independently propose a two-stage route for receptor action. Independently of Ariens he proposed that the binding of a ligand to a receptor produced a stimulus (S) that was related to tissue response. The magnitude of the stimulus depends on both the affinity of the ligand for the receptor and its efficacy (e).

Figure A pictorial representation of the variation of dose–response curves with the value of a. Values for a between 1 and 0 correspond to drugs that act as partial agonists, the degree of partial agonism depending on the value of a.

As a result, Clark’s equation (8.19) was modified to:

Equation (8.23) shows that ligands with an e value of zero will have no biological response. Consequently, full antagonists will have an e value of zero. To obtain a positive response e must have a positive value and so agonists and partial agonists will have positive e values. Moreover, the higher the positive value, the greater the response (Fig.) and the lower the dose of agonist [D] required to achieve the maximum response. This means that agonists with a high efficacy will produce a maximum response even though they do not occupy all of the available receptor sites. Unoccupied receptors are known as spare receptors. Their presence increases the sensitivity of a receptor to other ligands. It is now known that cells can contain several thousand receptors of a particular type. This number can increase (upregulation) or decrease (downregulation). These changes may be brought about by both pathological and physiological cell stimuli. They can affect drug response. For example, an increase (upregulation) in the number of receptors (RT) moves the drug response curve to a lower concentration, whilst a decrease will move it to a higher concentration (Fig.).

 

The effects on the dose–response curves of increasing e and RT

 

Stephenson observed that the magnitude of the response was not linearly related to the stimulus. This lead to a further modification of Clark’s equation to:

where f is a function known as the transducer function. The transducer function represents the properties of the signal transducer mechanism that links the signal from the ligand to the tissue response and is a characteristic of the responding tissue. As a result, the same ligand could have different transducer functions when it is bound to different tissues. This difference would explain why a ligand may act as an agonist in one tissue but as a partial agonist in a different tissue even though it is acting on the same receptor. Furthermore, differences in the transducer functions of different ligands acting on the same receptor in the same tissue would also explain why their relative potencies may be different. Potency depends on both the ligand–receptor complex and its efficacy. It is used to compare the relative effectiveness of different drugs and is defined as:

where e is the intrinsic efficacy, which is the efficacy per receptor, that is:

Since intrinsic efficacy is independent of the total number of available receptors, a drug with the same value for e in different tissues is likely to be acting on the same receptor in those tissues. Conversely, if the values are different, the drug is likely to be acting on different receptors in the different tissues.

 

Drugs for Treating Endoparasitic and Ectoparasitic Infestations

Adverse hygienic conditions favor human infestation with multicellular organisms (parasites). Skin and hair are colonization sites for arthropod ectoparasites, such as insects (lice, fleas) and arachnids (mites). Against these, insecticidal and arachnicidal agents, respectively, can be used. Endoparasites invade the intestines or even internal organs and are mostly members of

the phyla of flatworms and roundworms. They are combated with anthelmintics.

Antihelmintics. As shown in the table, the newer agents, praziquantel and mebendazole, are adequate for the treatment of diverse worm diseases. They are generally well tolerated, as are the other agents listed.

 

Insecticides. Whereas fleas can be effectively dealt with by disinfection of clothes and living quarters, lice and mites require the topical application of insecticides to the infested subject.

The following agents act mainly by interfering with the activation or inactivation of neural voltage-gated insect sodium channels.

 

Chlorphenothane (DDT) kills insects after absorption of a very low amount, e. g., via foot contact with sprayed surfaces (contact insecticide). The cause of death is nervous system damage and seizures. In humans DDT causes acute neurotoxicity only after absorption of very large amounts. DDT is chemically stable and is degraded in the environment and the body at extremely slow rates. As a highly lipophilic substance, it accumulates in fat tissues. Widespread use of DDT in pest control has led to its accumulation in food chains to alarming levels. For this reason, its use has now been banned in many countries.

 

Lindane is the active γ-isomer of hexachlorocyclohexane. It also exerts a neurotoxic action on insects (as well as humans). Irritation of skin or mucous membranes may occur after topical use. Lindane is active also against intradermal mites (Sarcoptes scabiei, causative agent of scabies), besides lice and fleas. Although it is more readily degraded than DDT, it should be used only as a second line agent with appropriate precautions. In the United Kingdom its use for head lice has been banned; in the United States it is not recommended in young children and is contraindicated in premature infants.

 

Permethrin, a synthetic pyrethroid, exhibits similar antiectoparasitic activity and may be the drug of choice owing to its slower cutaneous absorption, fast hydrolytic inactivation, and rapid renal elimination.

 

Antifungal agents

Fungal infections or mycoses may generally be divided into either superficial or systemic mycoses. Superficial mycoses affect the skin, nails, scalp and mucous membranes, while systemic mycoses affect internal tissues and organs. Since the middle of the twentieth century there has been an increase in both superficial and systemic mycoses. A significant degree of this increase is believed to be due to medical treatments, such as the use of antibiotics, radiotherapy, immunosuppressant drugs and steroids, which suppress a patient’s immune system. It is believed that this supression allows the fungal microorganisms to flourish. Fungal infections caused in this manner are referred to as opportunistic fungal infections. Opportunistic infections also occur in conditions such as AIDS where the immune system is suppressed by the disease.

 

Fungal microorganisms are believed to damage the cell membrane, leading to a loss of essential cellular components. This may result in inflammation of the infected tissue, which in some cases may be severe. Antifungal agents counter myoses by both fungistatic and fungicidal action. Fungistatic action occurs when a drug prevents the fungi reproducing, with the result that it dies out naturally, whilst fungicidal action kills the fungi. The suffixes -static and -cide are widely used to indicate these general types of action.

 

Fungal microorganisms differ from other microorganisms in that they consist of eukaryotic cells with a chitin cell wall. This means that their chemical structures and biochemistry are similar to those of humans. Consequently, it is more difficult to design drugs that would selectively target these fungi. However, there are some differences that can be utilised. For example, human cell membranes contain cholesterol but those of fungi contain ergosterol. A number of antifungal drugs are believed to act by blocking the biosynthesis of ergosterol in fungi.

 

Azoles

The azoles are a group of substituted imidazoles that exhibit fungistatic activity at nanomolar concentration and fungicidal activity at higher micromolar concentrations (Fig.). They are active against most fungi that infect the skin and mucous membrane. Azoles are also active against some systemic fungal infections, bacteria, protozoa and helminthic species.

(a) Examples of the structures of some active 1,3-diazoles. Note the common structural features. (b) Examples of azoles based on 1,2,4-triazole ring systems

 

An outline of the biosynthesis of ergosterol

In common with most drugs, the azoles are believed to act at a number of different sites, all of which contribute to their fungicidal action. However, their main point of action is believed to be the inhibition of some of the cytochrome P-450 oxidases found in the membranes of the microorganisms. In particular, azoles have been linked to inhibition of the enzyme 14a-sterol demethylase (P-450DM), which is essential for the biosynthesis of ergosterol, the main sterol found in the fungal cell membranes (Fig.). It is believed that nitrogen at position 3 of the imidazole rings (Fig. a) and nitrogen at position 4 of the triazole rings (Fig. b) bind to the iron of the haem units found in the enzyme, thereby blocking the action of the enzyme. This appears to lead to an accumulation of 14a-methylated sterols such as lanosterol in the membrane, which is thought to increase the membrane’s permeability, allowing essential cellular contents to leak causing irreversible cell damage and death. However, the precise details of the mode of action of azoles have yet to be fully elucidated. Azoles also inhibit the P-450 oxidases found in mammalian steroid biosynthesis, but in mammals much higher concentrations than those necessary for inhibition of the fungal sterol 14a-demethylases are usually required. Structure–action studies have shown that a weakly basic imidazole or 1,2,4-triazole rings substituted only at the N-1 position are essential for activity. The substituent must be lipophilic in character and usually contains one or more five- or six-membered ring systems, some of which may be attached by an ether, secondary amine or thioether group to

the carbon chain. The more potent compounds have two or three aromatic substituents, which in the more potent compounds are singly or multi-chlorinated or -fluorinated at positions 2, 4 and 6. These non-polar structures give the compounds a high degree of lipophilicity, and hence membrane solubility.

 

Allylamines and related compounds

Allylamines are synthetic derivatives of 3-aminopropene (Fig. ) developed from naftifine. They are weak bases, their hydrochlorides being only slightly soluble in water. The allylamine group appears to be essential for activity.  Allylamines are believed to act by inhibiting squalene epoxidase, the enzyme for the squalene epoxidation stage in the biosynthesis of ergosterol in the fungal membrane (Fig.). This leads to an increase in squalene concentration in the membrane with subsequent loss of membrane integrity, which allows loss of cell contents to occur. Tolnaftate, although it is not an allyl amine, appears to act in a similar fashion. However, allylamines do not appear to significantly inhibit the mammalian cholesterol biosynthesis.

Examples of the structures of allylamines. Naftidine and terbinafine are used as fungicides to treat deratophytes and filamentous fungi but only have a fungistatic action against pathogenic yeasts

 

Phenols

There are numerous phenolic antifungal agents (Fig.). They are believed to destroy sections of the cell membrane, which results in the loss of the cellular components and the death of the cell. The mechanism by which this destruction occurs is not known. Ciclopirox is not a phenol but appears to have a similar action. However, at low concentrations it has been shown to block the movement of amino acids into susceptible fungal cells.

 

Examples of phenolic compounds used as antifungal agents

 

Antibacterial antifungal agents

A number of antibiotics are important antifungal agents. They are mainly polyenes such as amphopterin B, nystatin and natamycin (Fig.). However, the antibiotic griseofulvin (Fig.d) is also active. The smaller polyenes (26-membered ring) exhibit both fungistatic and fungicidal action at the same concentration. In contrast, the larger ring polyenes (38-membered ring) show fungistatic action at lower concentrations and fungicidal action at higher concentrations. This indicates some differences in their mode of action. All the polyenes are believed to act by binding to the cell membrane, causing leakage of the cytoplasmic contents and cell lysis. It is thought that amphopterin B binds to the ergosterol found in the cell membranes of microfungi to form a transmembrane channel that allows the leakage of the cell contents. Unfortunately, it also appears to act in the same way with the cholesterol found in human cell membranes, which accounts for its toxic side effects in humans when administered by parenteral routes. The poor water solubility of polyenes means that they are difficult to administer by parenteral routes. However, amphopterin B may be administered parentally using micelle formulations. Consequently, their difficulty of administration and often unpleasant side effects results in polyenes being used mainly in topical preparations.

 

Griseofulvin is a fungistatic agent. It acts by preventing the infestation of new tissue as that tissue is formed. This is a slow process and so its use is only successful if the patient sticks rigidly to the prescribed drug regimen. Griseofulvin is used to treat systemic infections and has few side effects. It has a poor water solubility and so its oral adsorption and hence its effectiveness will depend on how the dose is formulated.

(a) Amphopterin B, first isolated from Streptomyces nodosus by Gold et al. (b) Nystatin, first isolated from Streptomyces noursei by Hazen and Brown. (c) Natamycin, first isolated from Streptomyces natalensis by Struyk et al. (d) Griseofulvin,first isolated from Penicillium griseofulvium by Oxford et al.

 

Chemotherapy of Viral Infections

Viruses essentially consist of genetic material (nucleic acids) and a capsular envelope made up of proteins, often with a coat of a phospholipid (PL) bilayer with embedded proteins. They lack a metabolic system and depend on the infected cell for their growth and replication. Targeted therapeutic suppression of viral replication requires selective inhibition of those metabolic processes that specifically serve viral replication in infected cells.

 

Viral replication as exemplified by herpes simplex viruses (A)

1. The viral particle attaches to the host cell membrane (adsorption) via envelope glycoproteins that make contact with specific structures of the cell membrane.

 

2. The viral coat fuses with the plasmalemma of host cells and the nucleocapsid (nucleic acid plus capsule) enters the cell interior (penetration).

 

3. The capsule opens (“uncoating”) near the nuclear pores and viral DNA moves into the cell nucleus. The genetic material of the virus can now direct the cell’s metabolic system.

 

4a. Nucleic acid synthesis: The genetic material (DNA in this instance) is replicated and RNA is produced for the purpose of protein synthesis.

 

4b. The proteins are used as “viral enzymes” catalyzing viral multiplication (e. g., DNA polymerase and thymidine kinase), as capsomers, or as coat components, or are incorporated into the host cell membrane.

 

5. Individual components are assembled into new virus particles (maturation).

6. Release of daughter viruses results in spread of virus inside and outside the organism. With herpesviruses, replication entails host cell destruction and development of disease symptoms.

 

Antiviral mechanisms (A)

The organism can disrupt viral replication with the aid of cytotoxic T-lymphocytes that recognize and destroy virus-producing cells (presenting viral proteins on their surface) or by means of antibodies that bind to and inactivate extracellular virus particles. Vaccinations are

designed to activate specific immune defenses.

 

Interferons (IFN) are glycoproteins that, among other products, are released from virus-infected cells. In neighboring cells, interferon stimulates the production of “antiviral proteins.” These inhibit the synthesis of viral proteins by (preferential) destruction of viral DNA or by suppressing its translation. Interferons are not directed against a specific virus, but have a broad spectrum of antiviral action that is, however, species-specific. Thus, interferon for use in humans must be obtained from cells of human origin, such as leukocytes (IFN-α), fibroblasts (IFN-β), or lymphocytes (IFN-γ). Interferons are used in the treatment of certain viral diseases, as well as malignant neoplasias and autoimmune diseases; e. g., IFN-α for the treatment of chronic hepatitis C and hairy cell leukemia; and IFN-β in severe herpes virus infections and multiple sclerosis.

 

Virustatic antimetabolites are “false” DNA building blocks (B) or nucleosides. A nucleoside (e. g., thymidine) consists of a nucleobase (e. g., thymine) and the sugar deoxyribose. In antimetabolites, one of the components is defective. In the body, the abnormal nucleosides undergo bioactivation by attachment of three phosphate residues.

 

Idoxuridine and congeners are incorporated into DNA with deleterious results. This also applies to the synthesis of human DNA. Therefore, idoxuridine and analogues are suitable only for topical use (e. g., in herpes simplex keratitis).

 

Chemotherapy of Malignant Tumors

A tumor (neoplasm) consists of cells that proliferate independently of the body’s inherent “building plan.” A malignant tumor (cancer) is present when the tumor tissue destructively invades healthy surrounding tissue or when dislodged tumor cells form secondary tumors (metastases) in other organs. A cure requires the elimination of all malignant cells (curative therapy). When this is not possible, attempts can be made to slow tumor growth and thereby prolong the patient’s life or improve quality of life (palliative therapy). Chemotherapy is faced

with the problem that the malignant cells are endogenous and almost lacking in specific metabolic properties.

 

Cytostatics (A)

Cytostatiscs are cytotoxic substances that particularly affect proliferating or dividing (mitotic) cells. Rapidly dividing malignant cells are preferentially injured. Damage to mitotic processes not only retards tumor growth but also may initiate apoptosis (programmed cell death). Tissues with a low mitosis rate are largely unaffected; likewise, most healthy tissues. This, however, also applies to malignant tumors consisting of slowly dividing differentiated cells. Tissues that have a physiologically high mitosis rate are bound to be affected by cytostatic therapy. Thus, typical adverse effects occur. Loss of hair results from injury to hair follicles; gastrointestinal disturbances, such as diarrhea, from inadequate replacement of enterocytes whose lifespan is limited to a few days; nausea and vomiting from stimulation of area postrema chemoreceptors; and lowered resistance to infection from weakening of the immune system. In addition, cytostatics cause bone marrow depression. Resupply of blood cells depends on the mitotic activity of bone marrow stem and daughter cells. When myeloid

proliferation is arrested, the short-lived granulocytes are the first to be affected (neutropenia), then blood platelets (thrombopenia) and, finally, the more long-lived erythrocytes (anemia). Infertility is caused by suppression of spermatogenesis or follicle maturation. Most cytostatics disrupt DNA metabolism. This entails the risk of a potential genomic alteration in healthy cells (mutagenic effect). Conceivably, the latter accounts for the occurrence of leukemias several years after cytostatic therapy (carcinogenic effect). Furthermore, congenital malformations are to be expected when cytostatics must be used during pregnancy (teratogenic effect). Cytostatics possess different mechanisms of action.

 

Damage to themitotic spindle (B)

The contractile proteins of the spindle apparatus must draw apart the replicated chromosomes before the cell can divide. This process is prevented by the so-called spindle poisons that arrest mitosis at metaphase by disrupting the assembly into spindle threads of microtubuli. These consist of the proteins α-and β-tubulin. Surplus tubuli are broken down, enabling the tubulin subunits to be recycled. The vinca alkaloids, vincristine and vinblastine (from the periwinkle plant, Vinca rosea), inhibit the polymerization of tubulin subunits into microtubuli. Damage to the nervous systemis a predicted adverse effect arising from injury to microtubule-operated axonal transport mechanisms.

 

Paclitaxel, from the bark of the pacific yew (Taxus brevifolia), inhibits disassembly of microtubules and induces formation of atypical ones, and thus impedes the reassemblage of

tubulins into properly functioning microtubules. Docetaxel is a semisynthetic derivative.

 

Inhibition of DNA and RNA synthesis (A)

Mitosis is preceded by replication of chromosomes (DNA synthesis) and increased protein synthesis (RNA synthesis). Existing DNA (gray) serves as a template for the synthesis of new (blue) DNA or RNA. De-novo synthesis may be inhibited by the following mechanisms.

 

Damage to the template (1)

Alkylating cytostatics

Alkylating cytostatics are reactive compounds that transfer alkyl residues into a covalent bond with DNA. For instance, mechlorethamine (nitrogen mustard) is able to cross-link double stranded DNA on giving off its chlorine atoms. Correct reading of genetic information is thereby rendered impossible. Other alkylating agents are chlorambucil, melphalan, thio-TEPA, cyclophosphamide, ifosfamide, lomustine, and busulfan. Specific adverse reactions include irreversible pulmonary fibrosis due to busulfan and hemorrhagic cystitis caused by the cyclophosphamide metabolite acrolein (preventable by the uro-protectant mesna = sodium 2-mercaptoethanesulfonate).

 

The platinum-containing compounds cisplatin and carboplatin release platinum, which binds to DNA. Cystostatic antibiotics insert themselves into the DNA double strand; this may lead to strand breakage (e. g., with bleomycin). The anthracycline antibiotics daunorubicin and adriamycin (doxorubicin)may induce cardiomyopathy. Bleomycin can also cause pulmonary fibrosis.

 

Induction of strand breakage may result from inhibition of topoisomerase. The epipodophyllotoxins etoposide and tenoposide interact with topoisomerase II, which functions to split, transpose, and reseal DNA strands; these agents cause strand breakage by inhibiting resealing. The “tecans” topotecan and irinotecan are derivatives of camptothecin from the fruits of a Chinese tree (Camptotheca acuminata). They inhibit topoisomerase I, which induces breaks in single-strand DNA.

 

Inhibition of nucleobase synthesis (2)

Tetrahydrofolic acid (THF) is required for the synthesis of both purine bases and thymidine. Formation of THF from folic acid involves dihydrofolate reductase (p. 274). The folate analogues aminopterin and methotrexate (amethopterin) inhibit enzyme activity.

Cellular stores of THF are depleted. The effect of these antimetabolites can be reversed by administration of folinic acid (5-formyl-THF, leucovorin, citrovorum factor). Hydroxyurea (hydroxycarbamide) inhibits ribonucleotide reductase that normally converts ribonucleotides

into deoxyribonucleotides subsequently used as DNA building blocks.

 

 

Incorporation of false building blocks (3)

Unnatural nucleobases (6-mercaptopurine; 5-fluorouracil) or nucleosides with incorrect sugars (cytarabine) act as antimetabolites. They inhibit DNA/RNA synthesis or lead to synthesis of missense nucleic acids. 6-Mercaptopurine results from biotransformation of the inactive precursor azathioprine. The uricostatic allopurinol inhibits the degradation of 6-mercaptopurine such that coadministration of the two drugs requires dose reduction of the latter.

 

Combination therapy

Cytostatics are frequently administered in complex therapeutic regimens designed to improve efficacy and tolerability of treatment.

 

Supportive therapy

Cancer chemotherapy can be supported by adjunctive medications. Thus, 5-HT3 serotonin receptor antagonists (e. g., ondansetron) afford effective protection against vomiting induced by highly emetogenic drugs such as cisplatin. Bone marrow depression can be counteracted by granulocyte and granulocyte/macrophage colony-stimulating factors (filgrastim and lenograstim and molgramostim, respectively).

 

Immunomodulation

The control of disease by immunologic means has two objectives: the development of immunity and the avoidance of undesired immune reactions. Immunostimulation in a drug-induced immunosuppression model and immunosuppression in an experimental hyperreactivity model by the same preparation can be said to be true immunomodulation. Immunomodulators are biological response modifi ers (BRMs), used to treat cancer, which exert their antitumor effects by improving host defense mechanisms against the tumor. They have a direct antiproliferative effect on tumor cells and also enhance the ability of the host to tolerate damage by toxic chemicals that may be used to destroy the cancer. Modulation of immunity was previously attempted with glucocorticoids and cytotoxic drugs such as cyclophosphamide. It is now recognized that immunomodulatory therapy could provide an alternative to conventional chemotherapy for a variety of diseased conditions, especially when the host’s defense mechanisms have to be activated under the conditions of impaired immune responsiveness or when a selective immunosuppression has to be induced in situations such as infl ammatory diseases, autoimmune disorders, and organ/bone marrow transplantation. All three classes of immunomodulators—biologicals, chemical, and cytokines—will continue to play a major role in advancing and improving the quality of treatment of several human as well as animal diseases. There is need for further research to better understand the biochemical mechanisms involved in immunoregulation to maximize the benefi ts of chemical immunomodulators as single agents or adjuvants in cancer therapy.

 

Ethnopharmacology and Botanical Immunomodulators

There are two major ways of bioprospecting natural products for investigation. The first is the classical method, which relies on phytochemical factors, serendipity, and random screening approaches. The second method uses traditional knowledge and practices as the drug discovery engine. Known as the ethnopharmacology approach, this method is time and cost-effective and may lead to better success than routine random screening. Various ethnopharmacological agents are under investigation as immunomodulators. Traditional Chinese medicine, Japanese Kampo, Indian Ayurveda, and such are becoming important bioprospecting tools. Ayurveda gives a separate class of immunomodulating botanicals named Rasayanas. Ayurveda, one of the most ancient and yet living traditions practiced idely in India, Sri Lanka, and other countries, has a sound philosophical and experiential basis. India has about 45,000 plant species; medicinal properties have been assigned to many to several thousands. Ayurveda has detailed descriptions of over 700 herbs and 400,000-registered Ayurvedic practitioners routinely prescribe them, particularly for treatment of chronic disease conditions. Considerable research on pharmacognosy, chemistry, pharmacology, and clinical therapeutics has been carried out and the Ayurvedic database has detailed descriptions of over 700 medicinal plants. Rasayanas are nontoxic herbal preparations or individual herbs used to rejuvenate or attain the complete potential of a healthy or diseased person in order to prevent diseases and degenerative changes that lead to disease. Pharmacodynamic studies on Rasayana botanicals have suggested many possible mechanisms, such as nonspecific and specific immunostimulation, free-radical quenching, cellular detoxification, cell proliferation, and cell repair. Ayurveda (with particular reference to botanicals) may play an important role in modern health care, particularly where satisfactory treatment is not available. There is a need to evaluate the potential of Ayurvedic remedies as adjuvant to counteract side effects of modern therapy and compare the cost-effectiveness of certain therapies vis-à-vis modern therapeutic schedules.

 

Adaptogens or Adjustive Medicine

Most of the synthetic chemotherapeutic agents available today are immunosuppressants, are cytotoxic, and exert a variety of side effects. N. V. Lazarev, who developed the concept of a state of nonspecifically increased resistance of an organism (SNIR), laid down the theoretical basis for separation of a new group of medicinal substances. The medicinal substances causing SNIR were named adaptogens. Generally, adaptogens are those drugs that enable one to withstand the stress and strain of life, impart immunity to give protection against diseases, postpone aging, and improve vigor, vitality, and longevity. The concept is also referred to as adjustive medicine. The concept of adjustive remedies has been difficult to prove experimentally. A bifunctional information exchange network between the nervous and immune systems is established by specific receptors for humoral substances on cells of nervous and immune systems. In particular, neuroregulators (neurotransmitters and neuromodulators) can modulate specific immune system function(s), and immunoregulators (immunomodulators) can modulate specific nervous system function(s). Acute and chronic inflammatory processes, malignancy, and immunological reactions stimulate the synthesis and release of immunomodulators in various cell systems.

 

Botanicals with Adaptogenic Activity

Mistletoe Lectin

Defined nontoxic doses of the galactoside-specific mistletoe lectin (mistletoe lectin-I, a constituent of clinically approved plant extract) have immunomodulatory potencies. The obvious ability of certain lectins to activate nonspecific mechanisms supports the assumption that lectin-carbohydrate interactions may induce clinically beneficial immunomodulation. Randomized multicenter trials are being performed to evaluate the ability of complementary mistletoe lectin-I treatment to reduce the rate of tumor recurrences and metastases, to improve overall survival and the quality of life, and to exert immunoprotection in cancer patients under tumor-destructive therapy.

 

Achyranthes Bidentata

Achyranthes bidentata polysaccharide (ABP) root extract (25 to 100 mg/kg, day _1 to 7) could inhibit tumor growth (S-180) by 31 to 40%. A combination of cyclophosphamide and ABP increased the rate of tumor growth inhibition by 58%. ABP could potentiate LAK cell activity and increase the Con A–induced production of tumor necrosis factor (TNF-â) from murine spleenocytes. The S-180 cell membrane content of sialic acid was increased, and phospholipid decreased after ABP had acted on cells for 24 hours. Data suggest that the antitumor mechanism of ABP may be related to potentiation of host immunosurveillance mechanism and the changes in cell membrane features.

 

Chemoprotection and immunomodulation.

 

Chemoprotection

Modern cancer therapy produces substantial acute and chronic toxicity, which impairs quality of life and limits the effectiveness of treatment. Recent clinical and laboratory data suggest that repair of treatment-related injury is a multiphase and continuous process providing multiple opportunities for pharmacologic intervention. A host of agents (toxicity antagonists) are under development that modulate normal tissue response or interfere with mechanisms of toxicity. Although significant challenges remain, the routine application of such agents promises to reduce treatment-related morbidity substantially and potentially to allow treatment intensification in high-risk disease. The concept of site-specific inactivation of cytotoxic anticancer agents has been explored with numerous modalities. The goal of such chemoprotection is to improve the therapeutic ratio of an agent by selectively reducing its toxicity in non-tumor-bearing tissue, which is target for dose-limiting toxicity. Furthermore, a chemoprotectant cannot add new toxicities that might otherwise limit the administration of maximally tolerated doses of chemotherapeutic agent.

Drug Targets and Current Trends

Chemoprotection and cytoprotection are studied under preventive oncology and are interchangeable terms in cancer chemotherapy. Preventive oncology applies pharmacological agents to reverse, retard, or halt progression of neoplastic cells to invasive malignancy. Cancer chemoprevention is one of the newer approaches in the management of cancer. Epidemiological observations, preclinical animal pharmacology, knockout models, cancer cell lines, and clinical trials have shown the efficacy of this approach. Many drug targets are under clinical development; prostaglandin pathway, estrogen receptor modulation, gluthathione peroxidase inhibition, and immunomodulation appear promising. Celecoxib, tamoxifen, retinoids, rexinoids, selenium, tocopherols, and mofarotene are some of the promising leads and are in clinical development. New opportunities in clinical chemoprevention research include investigating chemopreventive effects of phytochemicals. Safer immunomodulating agents suitable for long-term therapy remain an unmet therapeutic need. Figure gives a schematic overview of cytoprotection and immunomodulation.

 

Botanical Immunomodulators as Chemoprotectants

Withania Somnifera Withania somnifera is an official drug mentioned in the Indian Herbal Pharmacopoeia and Ayurvedic Pharmacopoeia. Studies indicate that W. somnifera (ashwagandha) (WS) possesses anti-inflammatory, antitumor, antistress, antioxidant, immunomodulatory, hemopoietic, and rejuvenating properties. The chemistry of WS has been studied extensively, and over 35 chemical constituents have been identified, extracted, and isolated. The biologically active chemical constituents are alkaloids (isopelletierine, anaferine), steroidal lactones (withanolides, withaferins), saponins containing an additional acyl group (sitoindoside VII and VIII), and withanolides with a glucose at carbon 27(sitoindoside IX and X).

 

The suppressive effect of cyclophosphamide-induced toxicity by WS extracts was observed in mice. Administration of WS extracts signifi cantly reduced leucopenia induced by cyclophosphamide treatment, resulting in an increase in bone marrow cellularity. Administration of W. somnifera extract for 5 days along with cyclophosphamide (CTX) (1.5 mmol/kg body weight, i.p.) reduced the CTX-induced urotoxicity.31 Treatment of W. somnifera resulted in the enhancement of interferon gamma (IFN-gamma), interleukin-2 (IL-2), and granulocyte macrophage colony stimulating factor (GM-CSF), which were lowered by cyclophosphamide administration. Pharmacodyanmic studies reveal that the major activity of

W. somnifera may be due to the enhancement of cytokine production and stem cell proliferation

and its differentiation. These studies indicate that W. somnifera could reduce the cyclophosphamide toxicity and its usefulness in cancer chemotherapy.

 

The major activity of WS may be due to stimulation of stem cell proliferation, indicating the fact that WS could reduce cyclophosphamide toxicity and its usefulness in cancer chemotherapy. WS was also shown to prevent lipid peroxidation (LPO) in stress-induced animals, indicating its adjuvant as well as chemoprotectant activity. Glycowithanolides, consisting of equimolar concentrations of sitoindosides VII to X and withaferin A, isolated from the roots of WS were evaluated for protection in iron-induced hepatoxicity in rats. Ten days of oral administration of these active principles, in graded doses (10, 20, and 50 mg/kg), resulted in attenuation of hepatic lipid peroxidation (LPO) and the serum enzymes alanine aminotransferase, aspartate aminotransferase, and lactate dehydrogenase during iron-induced hepatoxicity.58 Antistress activity observed with W. somnifera will be an additional benefit, along with chemoprotectant activity.

 

Tinospora Cordifolia Tinospora cordifolia is widely used in ayurvedic medicines and is known for its immunomodulatory, antihepatotoxic, antistress, and antioxidant properties. It has been used in combination with other plant products to prepare a number of ayurvedic preparations. The chemistry has been studied extensively, and its chemical constituents can be broadly divided into alkaloids, diterpenoids, steroids, flavanoids, and lignans. Reviews have appeared on quaternary alkaloids and biotherapeutic diterpene glucosides of Tinospora species. Much of the work has been carried out on berberine, jatrorrhizine, tinosporaside, and columbin. Extracts of T. cordifolia (TC) have been shown to inhibit lipid peroxidation and superoxide and hydroxyl radicals in vitro. The extract was also found to reduce the toxic side effects of cyclophosphamide (25 mg/kg, 10 days) in the mice hematological system by free-radical formation as seen from total white cell count, bone marrow cellularity, and esterase-positive cells. The active principles of TC were found to possess anticomplementary and immunomodulatory activities. TC is reported for its various immunopharmacological activities (e.g., inhibition of C3-convertase of the classical complement pathway). Humoral and cell-mediated immunity were reported for cardioside, cardifolioside A, and cardiol and their activation was more pronounced with increasing incubation time.61 Extracts of T. cordifolia has been shown to inhibit lipid peroxidation and superoxide and hydroxyl radicals in vitro. The extract was also found to reduce the toxic side effects of cyclophosphamide (25 mg/kg, 10 days) in the mice hematological system by free-radical formation as seen from total white cell count, bone marrow cellularity, and esterase-positive cells.

 

Botanical Immunomodulators as Antitumor Agents

Plant products have contributed several novel compounds that possess promising antitumor activity. Crude extract of W. somnifera root has a strong tumoricidal and tumor growth  inhibitory activity. Withaferin A, an alkaloid isolated from the leaves, has been reported to show marked tumor inhibitory activity in vitro against cells derived from human carcinoma of nasopharynx and experimental mouse tumors. A single i.p. dose of withaferin A injection 24 or 48 hours after Ehrlich’s ascites tumor transplantation produced an immediate growth reduction in 3 to 80% of mice, followed by complete disappearance of tumor cells in the peritoneal cavity of the surviving mice with no signs of tumor development. However, withaferin A is toxic in mice, with an LD50 of 54 mg/kg body weight after an i.p. injection. An effective dose of crude extract was much higher (a cumulative dose of more than 10 g—750 mg/kg daily for 15 days) with less toxicity than reported doses of purified withaferin A and withanolide D, which exhibited toxic effects. Crude extract included a range of chemicals (e.g., a few flavanoids, several alkaloids, and other withanolides) in addition to withaferin A. In the ayurvedic system of treatment, dry powders or crude extract is used, and hence the effects observed may not be attributed to a single component. The rationale for this type of treatment is that the toxicity of an active component may be counteracted by another component, which may not have the desired therapeutic property. Resinous material from a methanol extract and orange-colored oil from a petroleum ether extract of Semecarpus anacardium Linn. F. have been found to possess antitumor properties in a P388 lymphocytic leukemia model. Acetylated oil of Semicarpus anacardium, which itself does not possess antitumor activity against experimental transplantable tumors, enhances the antitumor effect of anticancer drugs such as mitomycin-C, 6-mercaptopurine, and methotrexate when used in combination against P388 and S180 (ascites) tumor systems.

 

Extracts of T. cordifolia have been shown to inhibit lipid peroxidation and superoxide and hydroxyl radicals in vitro. The extracts were also found to reduce the toxic side effects of cyclophosphamide administration in mice. Moreover, their administration partially reduced elevated lipid peroxides in serum and liver as well as alkaline phosphatase and glutamine pyruvate transaminase thus indicating the value of Tinospora extracts in reducing the chemotoxicity induced by free radical–forming chemicals. Crude saponins obtained from shoots of Asparagus racemosus [asparagus crude saponins (ACSs,] were found to have antitumor activity. They inhibited the growth of human leukemia HL-60 cells in culture and macromolecular synthesis in a dose- and time-dependent manner. The ACS in the range 75 to 100 mg/mL was cytostatic; at concentrations greater than 200 mg/mL it was cytocidal to HL-60 cells. ACSs at 6 and 50 mg/mL inhibited the synthesis of DNA, RNA, and protein in HL-60 cells by 41, 5, and 4% or by 84, 68, and 59%, respectively. The inhibitory effect of ACSs on DNA synthesis was irreversible.