Herd immunity (also called herd effect, community immunity, population immunity, or social immunity) is a form of indirect protection from infectious disease that occurs when a large percentage of a population has become immune to an infection, thereby providing a measure of protection for individuals who are not immune.
In a population in which a large number of individuals are immune,
chains of infection are likely to be disrupted, which stops or slows the
spread of disease.
The greater the proportion of individuals in a community who are
immune, the smaller the probability that those who are not immune will
come into contact with an infectious individual.
Individual immunity can be gained by recovering from an infection or through vaccination.
Some individuals cannot become immune due to medical reasons and in
this group herd immunity is an important method of protection. Once a certain threshold has been reached, herd immunity gradually eliminates a disease from a population.
This elimination, if achieved worldwide, may result in the permanent
reduction in the number of infections to zero, called eradication. This method was used for the eradication of smallpox in 1977 and for the regional elimination of other diseases. Herd immunity does not apply to all diseases, just those that are contagious, meaning that they can be transmitted from one individual to another. Tetanus, for example, is infectious but not contagious, so herd immunity does not apply.
The term herd immunity was first used in 1923.
It was recognized as a naturally occurring phenomenon in the 1930s when
it was observed that after a significant number of children had become
immune to measles, the number of new infections temporarily decreased, including among susceptible children. Mass vaccination to induce herd immunity has since become common and proved successful in preventing the spread of many infectious diseases.
Opposition to vaccination has posed a challenge to herd immunity,
allowing preventable diseases to persist in or return to communities
that have inadequate vaccination rates.
Effects
Protection of those without immunity
Some individuals either cannot develop immunity after vaccination or for medical reasons cannot be vaccinated. Newborn infants are too young to receive many vaccines, either for safety reasons or because passive immunity renders the vaccine ineffective. Individuals who are immunodeficient due to HIV/AIDS, lymphoma, leukemia, bone marrow cancer, an impaired spleen, chemotherapy, or radiotherapy
may have lost any immunity that they previously had and vaccines may
not be of any use for them because of their immunodeficiency.
Vaccines are typically imperfect as some individuals' immune systems
may not generate an adequate immune response to vaccines to confer
long-term immunity, so a portion of those who are vaccinated may lack
immunity. Lastly, vaccine contraindications may prevent certain individuals from becoming immune.
In addition to not being immune, individuals in one of these groups may
be at a greater risk of developing complications from infection because
of their medical status, but they may still be protected if a large
enough percentage of the population is immune.
High levels of immunity in one age group can create herd immunity for other age groups. Vaccinating adults against pertussis
reduces pertussis incidence in infants too young to be vaccinated, who
are at the greatest risk of complications from the disease. This is especially important for close family members, who account for most of the transmissions to young infants.
In the same manner, children receiving vaccines against pneumococcus
reduces pneumococcal disease incidence among younger, unvaccinated
siblings. Vaccinating children against pneumococcus and rotavirus has had the effect of reducing pneumococcus- and rotavirus-attributable hospitalizations for older children and adults, who do not normally receive these vaccines. Influenza (flu) is more severe in the elderly than in younger age groups, but influenza vaccines lack effectiveness in this demographic due to a waning of the immune system with age.
The prioritization of school-age children for seasonal flu
immunization, which is more effective than vaccinating the elderly,
however, has shown to create a certain degree of protection for the
elderly.
For sexually transmitted infections (STIs), high levels of immunity in one sex induces herd immunity for both sexes.
Vaccines against STIs that are targeted at one sex result in
significant declines in STIs in both sexes if vaccine uptake in the
target sex is high. Herd immunity from female vaccination does not, however, extend to homosexual males.
If vaccine uptake among the target sex is low, then the other sex may
need to be immunized so that that sex can be sufficiently protected.
High-risk behaviors make eliminating STIs difficult since even though
most infections occur among individuals with moderate risk, the majority
of transmissions occur because of individuals who engage in high-risk
behaviors.
For these reasons, in certain populations it may be necessary to
immunize high-risk persons or individuals of both sexes to establish
herd immunity.
Evolutionary pressure
Herd immunity itself acts as an evolutionary pressure on certain viruses, influencing viral evolution
by encouraging the production of novel strains, in this case referred
to as escape mutants, that are able to "escape" from herd immunity and
spread more easily. At the molecular level, viruses escape from herd immunity through antigenic drift, which is when mutations accumulate in the portion of the viral genome that encodes for the virus's surface antigen, typically a protein of the virus capsid, producing a change in the viral epitope. Alternatively, the reassortment of separate viral genome segments, or antigenic shift, which is more common when there are more strains in circulation, can also produce new serotypes. When either of these occur, memory T cells no longer recognize the virus, so people are not immune to the dominant circulating strain. For both influenza and norovirus, epidemics temporarily induce herd immunity until a new dominant strain emerges, causing successive waves of epidemics. As this evolution poses a challenge to herd immunity, broadly neutralizing antibodies and "universal" vaccines that can provide protection beyond a specific serotype are in development.
Serotype replacement
Serotype replacement, or serotype shifting, may occur if the prevalence of a specific serotype declines due to high levels of immunity, allowing other serotypes to replace it. Initial vaccines against Streptococcus pneumoniae significantly reduced nasopharyngeal carriage of vaccine serotypes (VTs), including antibiotic-resistant types, only to be entirely offset by increased carriage of non-vaccine serotypes (NVTs). This did not result in a proportionate increase in disease incidence though since NVTs were less invasive than VTs. Since then, pneumococcal vaccines that provide protection from the emerging serotypes have been introduced and have successfully countered their emergence.
The possibility of future shifting remains, so further strategies to
deal with this include expansion of VT coverage and the development of
vaccines that use either killed whole-cells, which have more surface
antigens, or proteins present in multiple serotypes.
Eradication of diseases
If herd immunity has been established and maintained in a population
for a sufficient time, the disease is inevitably eliminated—no more
endemic transmissions occurs.
If elimination is achieved worldwide and the number of cases is
permanently reduced to zero, then a disease can be declared eradicated. Eradication can thus be considered the final effect or end-result of public health initiatives to control the spread of infectious disease.
The benefits of eradication include ending all morbidity and mortality
caused by the disease, financial savings for individuals, health care
providers, and governments, and enabling resources used to control the
disease to be used elsewhere. To date, two diseases have been eradicated using herd immunity and vaccination: rinderpest and smallpox. Eradication efforts that rely on herd immunity are currently underway for poliomyelitis, though civil unrest and distrust of modern medicine have made this difficult. Mandatory vaccination may be beneficial to eradication efforts if not enough people choose to get vaccinated.
Free riding
Herd immunity is vulnerable to the free rider problem.
Individuals who lack immunity, particularly those who choose not to
vaccinate, free ride off the herd immunity created by those who are
immune.
As the number of free riders in a population increases, outbreaks of
preventable diseases become more common and more severe due to loss of
herd immunity. Individuals may choose to free ride for a variety of reasons, including the perceived ineffectiveness of a vaccine, believing that the risks associated with vaccines are greater than those associated with infection, mistrust of vaccines or public health officials, bandwagoning or groupthinking, social norms or peer pressure, and religious beliefs.
Individuals are more likely to free ride if vaccination rates are high
enough so as to convince a person that he or she may not need to be
immune since a sufficient number of others already are.
Mechanism
Individuals who are immune to a disease act as a barrier in the
spread of disease, slowing or preventing the transmission of disease to
others. An individual's immunity can be acquired via a natural infection or through artificial means, such as vaccination. When a critical proportion of the population becomes immune, called the herd immunity threshold (HIT) or herd immunity level (HIL), the disease may no longer persist in the population, ceasing to be endemic. This threshold can be calculated by taking R0, the basic reproduction number,
or the average number of new infections caused by each case in an
entirely susceptible population that is homogeneous, or well-mixed,
meaning each individual can come into contact with every other
susceptible individual in the population, and multiplying it by S, the proportion of the population who are susceptible to infection:
S can be rewritten as (1 - p) because p is the proportion of the population that is immune and p + S equals one. Then, the equation can be rearranged to place p by itself as follows:
- → →
With p being by itself on the left side of the equation, it can now be written as pc
to represent the critical proportion of the population needed to become
immune to stop the transmission of disease, or the herd immunity
threshold. R0 functions as a measure of contagiousness, so low R0 values are associated with lower HITs, whereas higher R0s result in higher HITs. For example, the HIT for a disease with an R0 of 2 is theoretically only 50%, whereas with disease with an R0 of 10 the theoretical HIT is 90%.
These calculations assume that the entire population is susceptible,
meaning no individuals are immune to the disease. In reality, varying
proportions of the population are immune to any given disease at any
given time. To account for this, the effective reproductive number Re, also written as Rt, or the average number of infections caused at time t, can found by multiplying R0 by the fraction of the population that is still susceptible. When Re
is reduced to and sustained below 1, the number of cases occurring in
the population gradually decreases until the disease has been
eliminated.
If a population is immune to a disease in excess of that disease's HIT,
the number of cases reduces at a faster rate, outbreaks are even less
likely to happen, and outbreaks that occur are smaller than they would
be otherwise. If Re increases to above 1, then the disease is neither in a steady state nor decreasing in incidence but is actively spreading through the population and infecting a larger number of people than usual.
A second assumption in these calculations is that populations are
homogeneous, or well-mixed, meaning that every individual comes into
contact with every other individual, when in reality populations are
better described as social networks as individuals tend to cluster
together, remaining in relatively close contact with a limited number of
other individuals. In these networks, transmission only occurs between
those who are geographically or physically close to one another. The shape and size of a network is likely to alter a disease's HIT, making incidence either more or less common. In heterogeneous populations, R0
is now considered to be a measure of the number of cases generated by a
"typical" infectious person, which depends on how individuals within a
network interact with each other.
Interactions within networks are more common than between networks, in
which case the most highly connected networks transmit disease more
easily, resulting in a higher R0 and a higher HIT than would be required in a less connected network.
In networks that either opt not to become immune or are not immunized
sufficiently, diseases may persist despite existing in better-immunized
networks.
Boosts
Vaccination
The primary way to boost levels of immunity in a population is through vaccination. Vaccination is originally based on the observation that milkmaids exposed to cowpox were immune to smallpox, so the practice of inoculating people with the cowpox virus began as a way to prevent smallpox.
Well-developed vaccines provide protection in a far safer way than
natural infections, as vaccines generally do not cause the diseases they
protect against and severe adverse effects are significantly less
common than complications from natural infections. The immune system does not distinguish between natural infections and vaccines, forming an active response
to both, so immunity induced via vaccination is similar to what would
have occurred from contracting and recovering from the disease. To achieve herd immunity through vaccination, vaccine manufacturers aim to produce vaccines with low failure rates and policy makers aim to encourage their use.
After the successful introduction and widespread use of a vaccine,
sharp declines in the incidence of diseases it protects against can be
observed, necessarily decreasing the number of hospitalizations and
deaths caused by such diseases.
Assuming a vaccine is 100% effective, then the equation used for
calculating the herd immunity threshold can be used for calculating the
vaccination level needed to eliminate a disease, written as Vc. Vaccines are usually imperfect however, so the effectiveness, E, of a vaccine must be accounted for:
From this equation, it can be observed that if E is less than (1 − 1/R0), then it is impossible to eliminate a disease, even if the entire population is vaccinated. Similarly, waning vaccine-induced immunity, as occurs with acellular pertussis vaccines, requires higher levels of booster vaccination to sustain herd immunity.
If a disease has ceased to be endemic to a population, then natural
infections no longer contribute to a reduction in the fraction of the
population that is susceptible. Only vaccination contributes to this
reduction.
The relation between vaccine coverage and effectiveness and disease
incidence can be shown by subtracting the product of the effectiveness
of a vaccine and the proportion of the population that is vaccinated, pv, from the herd immunity threshold equation as follows:
It can be observed from this equation that, ceteris paribus,
any increase in either vaccine coverage or vaccine effectiveness,
including any increase in excess of a disease's HIT, further reduces the
number of cases of a disease. The rate of decline in cases depends on a disease's R0, with diseases with lower R0 values experiencing sharper declines. Vaccines usually have at least one contraindication
for a specific population for medical reasons, but if both
effectiveness and coverage are high enough herd immunity can protect
these individuals. Vaccine effectiveness is often, but not always, adversely affected by passive immunity,
so additional doses are recommended for some vaccines while others are
not administered until after an individual has lost his or her passive
immunity.
Passive immunity
Individual immunity can also be gained passively, in which antibodies to a pathogen are transferred from one individual to another. This can occur naturally, whereby maternal antibodies, primarily immunoglobulin G antibodies, are transferred across the placenta and in colostrum to fetuses and newborns, or artificially, by which antibodies from the serum or plasma of an immune individual are injected into a susceptible person.
Protection generated from passive immunity is immediate but wanes over
the course of weeks to months, so any contribution to herd immunity is
temporary.
For diseases that are especially severe among fetuses and newborns,
such as influenza and tetanus, pregnant women may be immunized in order
to transfer antibodies to the child.
In the same way, high-risk groups that are either more likely to
experience infection or are more likely to develop complications from
infection may receive antibody preparations to prevent these infections
or to reduce the severity of symptoms.
Cost–benefit analysis
Herd immunity is often accounted for when conducting cost–benefit analyses of vaccination programs. It is regarded as a positive externality
of high levels of immunity, producing an additional benefit of disease
reduction that would not occur had no herd immunity been generated in
the population.
Therefore, herd immunity's inclusion in cost–benefit analyses results
in more favorable cost-effectiveness or cost–benefit ratios and an
increase in the number of disease cases averted by vaccination.
Study designs done to estimate herd immunity's benefit include
recording disease incidence in households in which a member was
vaccinated, randomizing a population in a single geographic area to be
vaccinated or not, and observing disease incidence before and after a
vaccination program is introduced.
From these, it can be observed that disease incidence may decrease to a
level beyond what can be predicted from direct protection alone,
indicating that herd immunity contributed to the reduction. When serotype replacement is accounted for, it reduces the predicted benefits of vaccination.
History
The term herd immunity was first used in 1923 to refer to an entire population's immunity, in reference to research examining disease mortality in mouse populations with varying degrees of immunity.
Herd immunity was first recognized as a naturally occurring phenomenon
in the 1930s when A. W. Hedrich published research on the epidemiology of measles in Baltimore
and took notice that after many children had become immune to measles,
the number of new infections temporarily decreased, including among
susceptible children. In spite of this knowledge, efforts to control and eliminate measles were unsuccessful until mass vaccination using the measles vaccine began in the 1960s.
Mass vaccination, discussions of disease eradication, and cost–benefit
analyses of vaccination subsequently prompted more widespread use of the
term herd immunity. In the 1970s, the theorem used to calculated a disease's herd immunity threshold was developed. During the smallpox eradication campaign in the 1960s and 1970s, the practice of ring vaccination,
of which herd immunity is integral to, began as a way to immunize every
person in a "ring" around an infected individual to prevent outbreaks
from spreading.
Since the adoption of mass and ring vaccination, complexities and challenges to herd immunity have arisen.
Modeling of the spread of infectious disease originally made a number
of assumptions, namely that entire populations are susceptible and
well-mixed, which do not exist in reality, so more precise equations
have been developed.
In recent decades, it has been recognized that the dominant strain of a
microorganism in circulation may change due to herd immunity, either
because of herd immunity acting as an evolutionary pressure or because
herd immunity against one strain allowed another already-existing strain
to spread. Emerging or ongoing vaccine controversies
and various reasons for opposing vaccination have reduced or eliminated
herd immunity in certain communities, allowing preventable diseases to
persist in or return to these communities.