By Alex Elliot, BSc, and Joanna Ellis, PhD
Each year, increased hospitalisation rates and between 10,000 and 13,000 deaths (in an average AH3N2 year*) are attributable to influenza in the UK.1 Annual vaccination continues to be the main strategy for the control of influenza. Current influenza vaccines contain inactivated (killed) virus and new vaccines containing attenuated live viruses are undergoing clinical trials. The recent development of neuraminidase inhibitors provides a new tool for both the prevention and treatment of influenza infection.
Clinical features of influenza
Influenza is a highly infectious viral infection, which occurs mainly in the
winter months of October to March in the northern hemisphere. It is characterised
by a rapid onset with fever, chills, myalgia, headache and a non-productive
cough. The virus is transmitted by breathing in the tiny droplets from the breath
of infected people.
Influenza for most people is a mild illness, which is resolved in one to two
weeks. However, complications associated with influenza infection can occur
in both the upper and lower respiratory tract. Otitis media is a common complication
in children and conjunctivitis occurs in both adults and children. Serious respiratory
complications, such as bronchitis and pneumonia, which may be fatal, can occur.2
The potential for developing complications is higher in certain risk groups,
such as the elderly and individuals with chronic medical conditions.
Influenza diagnosis
Several different pathogens can produce respiratory illnesses with similar clinical
symptoms, making an accurate diagnosis of influenza by a physician difficult.
A number of laboratory tests are available to confirm the diagnosis of influenza.
These are based on either the detection of viral antigens, viral nucleic acid,
virally infected cells or infectious virus particles in respiratory secretions.
In addition, several rapid (less than 15 minutes to perform) near-patient tests
are being developed.3-5 The impact of such tests on the speed of influenza diagnosis
has yet to be assessed. Furthermore, attempts must still be made to isolate
virus for further detailed analysis and strain characterisation, for the purpose
of vaccine strain selection. Serological tests, which detect a rise in specific
antibody to the virus in a patients serum, are also performed where confirmation
of recent influenza infection is required.
The virus
Influenza viruses are divided into three types, designated A, B and C.6 Influenza
A viruses are further classified into subtypes based on differences in their
surface glycoproteins, haemagglutinin (HA) and neuraminidase (NA) (Figure 1).
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Figure 1: Diagrammatic representation
of an influenza virus particle |
To date, 15 HA and nine NA subtypes have been identified.7,8 All influenza A subtypes have been found in aquatic and domestic birds, but only a few subtypes have been found to infect mammals and humans. Two subtypes of influenza A are currently circulating in humans, H1N1 and H3N2. Influenza types B and C exclusively infect humans. Influenza A and B are responsible for epidemics of respiratory illness that occur almost annually and are often associated with increased rates of hospitalisation and death. Studies suggest that H3N2 infections are more severe than H1N1, and that influenza B is intermediate between the two.9 Influenza C viruses usually cause a mild or asymptomatic infection limited to the upper respiratory tract. Therefore, efforts to control the impact of influenza on public health are aimed at types A and B.
Evading immunity
The ability of influenza viruses to cause repeated infections is due to the
constantly changing structure of the virus. Both the HA and NA proteins are
antigenic, inducing the production of antibodies in the host. Spontaneous mutations
continually occur in HA and NA molecules, causing a gradual change in the virus.
This ongoing process occurs in both influenza A and B and is called antigenic
drift. The emergence, through drift, of new strains of influenza virus leads
to a reduction in the effectiveness of antibodies to previous infections, and
results in the ability of the virus to reinfect individuals and cause annual
epidemics of the disease. Worldwide epidemics, or pandemics, of influenza have
occurred at less frequent and irregular intervals. Pandemics are the consequence
of more radical changes brought about when genes from different strains or subtypes
of influenza A viruses infect the same host. The influenza viral genome is segmented,
making possible the interchange of genes from different viruses. This may result
in the emergence of a new influenza variant with a novel pairing of HA and NA
proteins, to which existing antibodies are ineffective.10 This process of genetic
reassortment, leading to a completely novel antigenic subtype in humans, is
termed antigenic shift.
Surveillance and selection of vaccine strains
As a result of the constantly changing make-up of influenza viruses, the World
Health Organisation (WHO) global influenza surveillance network monitors the
viruses causing outbreaks of influenza throughout the year in different parts
of the world. The network comprises four international centres located in London,
Atlanta, Melbourne and Tokyo, and 110 national laboratories in 83 countries.11
The aim of this surveillance is to collect, analyse and distribute information
on influenza activity, which will aid the prevention and control of influenza
infection and its complications. Three types of information are assessed. First,
epidemiological data, such as consultation rates with physicians for influenza-like
illness, illness levels among schoolchildren and death rates from respiratory
illnesses, are estimated. Secondly, antigenic and genetic characteristics of
the HA and NA proteins of viruses isolated from respiratory secretions in laboratories
are analysed. These are compared with viruses isolated from previous years and
those in the current vaccine. Thirdly, the ability of currently available vaccines
to evoke antibody responses in children, adults and the elderly to newly detected
strains is also analysed in clinical trials. All data from around the world
are collated by the WHO in Geneva, which, every year, recommends to vaccine
manufacturers which strains to include in the vaccine.
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Panel 1: Recommendations for influenza vaccine composition 2000/2001 |
Recommendations are made in February for vaccines to be used in the northern hemisphere the following year (Panel 1), and in September for use in the southern hemisphere.
Current influenza vaccines
Every year, new influenza vaccine has to be produced. The vaccine normally contains
three components: two subtypes of influenza A (H1N1 and H3N2) and one of influenza
B. The information on which strains have been dominant over the winter and any
evidence of the emergence and potential spread of new strains is carefully analysed.
The effectiveness of the vaccine will depend on how close the vaccine strains
match the influenza viruses that circulate in the following winter. Influenza
vaccines currently available in the UK contain highly purified inactivated (killed)
virus, grown in specific pathogen-free fertile hens eggs. Both split virion
and surface antigen vaccines (purified from chemically disrupted viruses) are
available, containing split particles, or HA and NA proteins, respectively.
The timing of the WHO recommendations is critical since it may take six to nine
months from when production begins in March to prepare and test the new vaccine
ready for the new influenza season. In Europe, the first step is the approval
of strains by the European Medicines Evaluation Agency in March. Standardisation
of the vaccine then takes place, in which potency and safety tests are performed.
The immunogenicity of the newly manufactured vaccine is then assessed in clinical
trials.
Immunisation policy
The aim of the Department of Health influenza immunisation policy is to reduce the level of morbidity and mortality due to influenza. This year, the recommendations for those to receive the vaccine have been extended to include immunisation of all people aged 65 and over,12 rather than 75 years and over as previously recommended (Panel 2).
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Panel 2: Recommended groups to receive influenza vaccination |
The decision follows an assessment showing that immunising the otherwise fit 65-74 year age group offers benefits in life expectancy as well as reductions in complications and hospital admissions. Despite efforts at targeting the vaccine, not all individuals in key target groups are receiving the vaccine.13,14 For the first time, a target of 70 per cent uptake in people aged 65 years and over is being set for health authorities, with the aim of a minimum 60 per cent uptake in this first year. The number of doses of influenza vaccine available to general practitioners in the UK will be increased from eight to 10 million.
New approaches to influenza vaccination
Inactivated influenza vaccine is highly effective in young adults. However,
efficacy may be suboptimal in children and the elderly. Certain individuals
are also allergic to egg proteins. Because of these limitations, efforts have
been directed towards either improving the efficacy of the current vaccine,
by the addition of new adjuvants together with intranasal administration, or
developing new types of vaccines. Other types of vaccine currently being studied
include:
Live attenuated vaccines An intranasal cold-adapted vaccine is in late clinical development in several countries.15 A donor virus, which has become attenuated by cold-adaptation, is mixed with a wild-type virus that contains the HA and NA segments against which protection is sought. Following reassortment of viral genes, the resulting vaccine strain contains an attenuated donor genome, but expresses the wild-type HA and NA surface proteins. Live attenuated vaccines have the advantage of stimulating stronger immune responses without causing illness and can be administered by spraying into the nose. Both live attenuated vaccines and recombinant vaccines (proteins produced by DNA technology) offer promise for the future and appear to be safe, with few severe reactions in vaccinated subjects, including very young infants.16
DNA vaccines The potential of immunisation with purified DNA as a powerful technique for inducing immune responses was first demonstrated in the early 1990s. The routes of inoculation used for influenza DNA immunisation include intramuscular, intradermal and delivery of DNA-coated gold beads directly to the skin. Although DNA vaccines have several advantages, such as no requirement for production of vaccine in eggs, the introduction of genetic information into mammalian hosts raises several safety concerns. These include the possible formation of anti-DNA antibodies and the potential for causing transformation from a normal cell to a cancer cell. Efficacy of DNA vaccination against influenza has been demonstrated in animals. Initial human trials are still in progress, so it is too early to predict if DNA vaccines may be used in human immunisation programmes.
Treatment of influenza infection
General treatment Influenza for most people is an unpleasant, but self-limiting,
disease. The main symptoms may last for up to seven days. Treatment for most
individuals is symptomatic and those affected are advised to stay at home, rest
and drink plenty of fluids.
Antiviral drugs
There is normally a good match between circulating viruses and the vaccine and
when this happens influenza vaccine has been shown to prevent illness in approximately
70-90 per cent of healthy individuals. This, however, leaves a window of vaccine
failures and at-risk patients where drug prophylaxis and treatment can provide
the protection needed.
Amantadine and rimantadine Until last year, the anti-matrix protein drugs amantadine (Symmetrel), and its close derivative rimantadine (Flumavine) were the only antiviral compounds licensed for the treatment of influenza infections.
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| Figure 2: Amantadine blocks the pore of the M2 ion channel located within the virus envelope preventing virus replication. (Figure from Action of adamantanamines by A. J. Hay in Seminars in Virology. Copyright © 1992 by Academic Press, reproduced by permission of the publisher) |
The target of both drugs is the M2 protein which forms an ion channel and facilitates the release of viral genetic material into the infected cell. Amantadine and rimantadine inhibit viral replication by blocking this channel (Figure 2).
The effectiveness of amantadine and rimantadine for both the prophylaxis and
treatment of influenza infections has been established through many studies
over the past 30 years. The benefits of prophylactic treatment have been proven
using experimental challenges in healthy individuals with influenza A.17 Vital
dose-response studies have helped determine the optimum dose for protection,
as both of the drugs have a low toxic-to-therapeutic ratio that makes dosing
vitally important.18 In naturally occurring influenza infections in adults and
children, amantadine and rimantadine have been shown to have an efficacy of
up to 90 per cent in preventing influenza illness.19,20 Within closed populations,
eg, boarding schools, nursing homes and prisons, the drugs have also been shown
to be effective in preventing illness, and, in some cases, reducing mortality.
21,22
Amantadine and rimantadine have proved equally effective in treating acute influenza
infections in healthy adults, children and the elderly.23-25 The drugs are equally
as effective as the annual vaccine and are able to prevent illness or reduce
influenza-like symptoms. However, it is thought that administration of amantadine
and rimantadine must be initiated within 48 hours of the onset of symptoms if
they are to be effective.
The above trials and studies prove that amantadine and rimantadine are potent
inhibitors of influenza and reduce disease, but their use in the UK has been
limited. The drugs are restricted in that they are only specific against influenza
A because their target protein, M2 is present only in influenza A viruses. There
has been concern about the side effects associated with treatment. It has been
reported that these occur in approximately 6 per cent of patients, more often
in the elderly, and can consist of neurological reactions, including light-headedness
and an inability to concentrate, and gastrointestinal complaints.26, 27 Due
to the lower incidence of neurological side effects associated with rimantadine
treatment, it is now perceived to be the drug of choice in some countries, although
it can still cause gastrointestinal complaints, nausea and vomiting.18 In the
UK, amantadine has been rebranded (Lysovir) and the recommended dosage of the
drug has been reduced. The overall efficacy has been retained at this lower
dose but the occurrence of side effects has been reduced.
In vitro and in vivo studies have shown that cross-resistant viruses to both
drugs can be generated very rapidly.26 Furthermore, transmission of resistant
viruses between humans has been observed in families undergoing prophylactic
drug treatment28 and can cause typical influenza.29
In general, there seems to be a lack of awareness of the potential of these
drugs for preventing or limiting influenza A infections. With the advent of
novel inhibitors, amantadine and rimantadine may still play an important role
in the control of influenza through combined drug therapies.
Neuraminidase inhibitors The NA molecule located on the surface of the influenza virus particle (Figure 1) is an enzyme that is essential for virus replication and infectivity. The enzyme cleaves terminal sialic acid residues (the receptor molecules for the viral haemagglutinin) from neighbouring glycoconjugates. The active site of the molecule is highly conserved in NA species of both influenza A and B viruses, making it an ideal target for antiviral therapy.
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| Figure 3: Diagrammatic representation of NI compounds blocking the active site of the NA molecule. Green lines represent bonds formed between different groups on the drugs and amino acids within the site. Zanamivir and oseltamivir bind to the site more readily than sialic acid through a guanidine and hydrophobic group, respectively, increasing their antiviral potency (Adapted from Disarming flu viruses, by W. Graeme Laver, Norbert Bischofberger and Robert G. Webster. Copyright © 1998 by Scientific American, Inc.) |
The principle behind neuraminidase inhibitors (NI) is to provide a synthetic
sialic acid residue that will block the active site of the enzyme and prevent
it from functioning (Figure 3). There has been much research carried out in
this field which has encouraged the development of new sialic acid derivatives
with potent antiviral properties. One of these compounds, GG167 (zanamivir),
was discovered in 199330 and has been developed by Glaxo Wellcome under licence
from the Australian company Biota Holdings. Phase I and II clinical trials proved
its potency and phase III trials have proved its efficacy and safety in human
subjects.
Results from randomised, double-blind placebo-controlled studies of large groups
of subjects in both the northern31-35 and the southern36 hemispheres have demonstrated
the clinical usefulness of the drug. The main conclusions were that in subjects
treated with zanamivir, symptoms of headache, sore throat, fever, cough, weakness
and muscle ache were alleviated faster than in placebo groups, a difference
of one to 2.5 days. However, for the drug to be clinically effective it had
to be administered less than 30-40 hours after the onset of symptoms. The incidence
of complications and use of antibiotics were also reported to be lower in treated
groups. Side effects were described in approximately 3 per cent of subjects,
the most common being sinusitis, diarrhoea and nausea, but this was comparable
to placebo groups. Zanamivir is applied topically to the respiratory tract,
via inhalation using a Diskhaler,33 but it has been reported that this can cause
bronchospasm and/or a decline in lung function in some patients with underlying
respiratory disease.
The application for licensing zanamivir, under the trade name Relenza, was initially
turned down by the US Food and Drug Administration (FDA) on the grounds that
the US trials did not produce as good results as other trials around the world.37
The FDA was eventually persuaded after further trials and more conclusive results
relating to the clinical efficacy and safety of the drug. Zanamivir was licensed
in the US in July, 1999, and in the UK in September, 1999. Controversially,
the National Institute for Clinical Excellence (NICE) in the UK, after agreeing
to set-up a fast track assessment of the drug to enable its use,
recommended that zanamivir should not be prescribed in the 1999/2000 winter
season. The NICE stated that further studies needed to be completed to fully
assess clinical benefits, especially in at-risk groups.38
Another promising NI, oseltamivir was first described in 199739 and was licensed
in the US in late 1999, under the trade name Tamiflu. The main advantages oseltamivir
has over zanamivir is that its bioavailability is much greater and it can be
taken orally. The efficacy and safety of oseltamivir has been proven and the
drug is effective at reducing the duration and severity of illness.40-44 Oseltamivir
significantly reduced the duration of illness in infected patients by 25 per
cent. Where treatment was initiated within 24 hours of the onset of symptoms
this reduction was 37 per cent. It was reported that the overall health of treated
patients was improved in respect of symptoms, activity and sleep quality, and
secondary complications, including sinusitis, bronchitis and pneumonia, were
also reduced. Overall conclusions show that oseltamivir is well tolerated and
treatment reduces the severity and duration of acute influenza infection.
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Panel 3: Useful links containing influenza-related information
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Previous studies of amantadine and rimantadine showed how resistance could
be generated in patients undergoing treatment.23 There is similar concern with
NIs. With the predicted demand for the drugs, will the isolation of resistant
viruses increase? In vitro studies have demonstrated that growing the virus
repeatedly in the presence of drug does lead to resistance but to date there
have been few resistant viruses isolated from clinical cases. As the demand
for NIs increases, it will be important to monitor the generation and genetic
composition of resistant viruses.
The targeted use of NI drugs in prophylaxis, in special situations, would be
a beneficial alternative or addition to vaccination. When used prophylactically,
zanamivir has been shown to be highly effective in preventing infection with
clinical illness.34
New NI drugs are continually being developed. One such compound, RWJ-270201,
produced by Biocryst Pharmaceuticals, is currently undergoing phase II clinical
trials. Preliminary results from these early trials, suggest that human volunteers
infected with influenza showed significant reductions in virus titres after
treatment.
Further trials will be required to determine the full clinical effectiveness
of the drug before it appears on the market.
Antiviral therapy has the potential to reduce the burden of influenza infection
on the general population. Effective treatment requires accurate diagnosis of
influenza infection within 48 hours of onset of symptoms. This is often very
difficult to achieve and therefore cost-effective use of drugs may rely upon
near-patient testing to confirm infection.
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Summary: Influenza prevention and treatment
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References
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