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Gene drive
A gene drive is a natural
process[1] and technology of
genetic engineering that
propagates a particular suite
of genes throughout a
population[2] by altering the
probability that a specific
allele will be transmitted to
offspring (instead of the
Mendelian 50% probability).
Gene drives can arise
through
a
variety
of
[3][4]
mechanisms.
They have
been proposed to provide an
effective
means
of
genetically
modifying
specific populations and
entire species.
The technique can employ adding, deleting, disrupting, or modifying genes.[5][6]
Proposed applications include exterminating insects that carry pathogens (notably mosquitoes that
transmit malaria, dengue, and zika pathogens), controlling invasive species, or eliminating herbicide
or pesticide resistance.[7][5][8][9]
As with any potentially powerful technique, gene drives can be misused in a variety of ways or induce
unintended consequences. For example, a gene drive intended to affect only a local population might
spread across an entire species. Gene drives used to eradicate populations of invasive species in their
non-native habitats may have consequences for the population of the species as a whole, even in its
native habitat. Any accidental return of individuals of the species to its original habitats, through
natural migration, environmental disruption (storms, floods, etc.), accidental human transportation,
or purposeful relocation, could unintentionally drive the species to extinction if the relocated
individuals carried harmful gene drives.[10]
Gene drives can be built from many naturally occurring selfish genetic elements that use a variety of
molecular mechanisms.[3] These naturally occurring mechanisms induce similar segregation
distortion in the wild, arising when alleles evolve molecular mechanisms that give them a
transmission chance greater than the normal 50%.
Most gene drives have been developed in insects, notably mosquitoes, as a way to control insect-borne
pathogens. Recent developments designed gene drives directly in viruses, notably herpesviruses.
These viral gene drives can propagate a modification into the population of viruses, and aim to reduce
the infectivity of the virus.[11][12]
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Contents
Mechanism
Molecular mechanisms
Spreading in the population
Gene drive in viruses
Technical limitations
Issues
Bioethics concerns
History
Funding
Control strategies
CRISPR
Applications
Disease vector species
Invasive species control
Wild animal welfare
See also
References
Further reading
External links
Mechanism
In sexually-reproducing species, most genes are present in two copies (which can be the same or
different alleles), either one of which has a 50% chance of passing to a descendant. By biasing the
inheritance of particular altered genes, synthetic gene drives could spread alterations through a
population.[5][6]
Molecular mechanisms
At the molecular level, an endonuclease gene drive works by cutting a chromosome at a specific site
that does not encode the drive, inducing the cell to repair the damage by copying the drive sequence
onto the damaged chromosome. The cell then has two copies of the drive sequence. The method
derives from genome editing techniques and relies on the fact that double strand breaks are most
frequently repaired by homologous recombination, (in the presence of a template), rather than nonhomologous end joining. To achieve this behavior, endonuclease gene drives consist of two nested
elements:
either a homing endonuclease or a RNA-guided endonuclease (e.g., Cas9 or Cas12a[13]) and its
guide RNA, that cuts the target sequence in recipient cells
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a template
sequence used
by the DNA
repair machinery
after the target
sequence is cut.
To achieve the
self-propagating
nature of gene
drives, this repair
template
contains at least
the
endonuclease
sequence.
Because the
template must be
used to repair a
double-strand
break at the
cutting site, its
Molecular mechanism of gene drive.
sides are
homologous to
the sequences
that are adjacent to the cutting site in the host genome. By targeting the gene drive to a gene
coding sequence, this gene will be inactivated; additional sequences can be introduced in the
gene drive to encode new functions.
As a result, the gene drive insertion in the genome will re-occur in each organism that inherits one
copy of the modification and one copy of the wild-type gene. If the gene drive is already present in the
egg cell (e.g. when received from one parent), all the gametes of the individual will carry the gene
drive (instead of 50% in the case of a normal gene).[5]
Spreading in the population
Since it can never more than double in frequency with each generation, a gene drive introduced in a
single individual typically requires dozens of generations to affect a substantial fraction of a
population. Alternatively, releasing drive-containing organisms in sufficient numbers can affect the
rest within a few generations; for instance, by introducing it in every thousandth individual, it takes
only 12–15 generations to be present in all individuals.[14] Whether a gene drive will ultimately
become fixed in a population and at which speed depends on its effect on individual fitness, on the
rate of allele conversion, and on the population structure. In a well mixed population and with
realistic allele conversion frequencies (≈90%), population genetics predicts that gene drives get fixed
for selection coefficient smaller than 0.3;[14] in other words, gene drives can be used to spread
modifications as long as reproductive success is not reduced by more than 30%. This is in contrast
with normal genes, which can only spread across large populations if they increase fitness.
Gene drive in viruses
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Because the strategy usually relies on the simultaneous presence of an unmodified and a gene drive
allele in the same cell nucleus, it had generally been assumed that a gene drive could only be
engineered in sexually reproducing organisms, excluding bacteria and viruses. However, during a
viral infection, viruses can accumulate hundreds or thousand genome copies in infected cells. Beside,
cells are frequently co-infected by multiple virions and recombination between viral genomes is a
well-known and widespread source of diversity for many viruses. In particular, herpesviruses are
nuclear-replicating DNA viruses with large double-stranded DNA genomes and frequently undergo
homologous recombination during their replication cycle.
These properties have enabled the design of a gene drive strategy that doesn't involve sexual
reproduction, but relies on co-infection of a given cell by a naturally occurring and an engineered
virus. Upon co-infection, the unmodified genome is cut and repaired by homologous recombination,
producing new gene drive viruses that can progressively replace the naturally occurring population. In
cell culture experiments, it was shown that a viral gene drive can spread into the viral population and
strongly reduce the infectivity of the virus, which opens novel therapeutic strategies against
herpesviruses.[11]
Technical limitations
Because gene drives propagate by replacing other alleles that contain a cutting site and the
corresponding homologies, their application has been mostly limited to sexually reproducing species
(because they are diploid or polyploid and alleles are mixed at each generation). As a side effect,
inbreeding could in principle be an escape mechanism, but the extent to which this can happen in
practice is difficult to evaluate.[15]
Due to the number of generations required for a gene drive to affect an entire population, the time to
universality varies according to the reproductive cycle of each species: it may require under a year for
some invertebrates, but centuries for organisms with years-long intervals between birth and sexual
maturity, such as humans.[16] Hence this technology is of most use in fast-reproducing species.
Effectiveness in real practice varies between techniques, especially by choice of germline promoter.
Lin and Potter 2016 (a) discloses the promoter technology homology assisted CRISPR knockin
(HACK) and Lin and Potter 2016 (b) demonstrates actual use, achieving a high proportion of altered
progeny from each altered Drosophila mother.[17]
Issues
Issues highlighted by researchers include:[18]
Mutations: A mutation could happen mid-drive, which has the potential to allow unwanted traits to
"ride along".
Escape: Cross-breeding or gene flow potentially allow a drive to move beyond its target
population.
Ecological impacts: Even when new traits' direct impact on a target is understood, the drive may
have side effects on the surroundings.
The Broad Institute of MIT and Harvard added gene drives to a list of uses of gene-editing technology
it doesn't think companies should pursue.[19]
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Bioethics concerns
Gene drives affect all future generations and represent the possibility of a larger change in a living
species than has been possible before.[20]
In December 2015, scientists of major world academies called for a moratorium on inheritable human
genome edits that would affect the germline, including those related to CRISPR-Cas9 technologies,[21]
but supported continued basic research and gene editing that would not affect future generations.[22]
In February 2016, British scientists were given permission by regulators to genetically modify human
embryos by using CRISPR-Cas9 and related techniques on condition that the embryos were destroyed
in seven days.[23][24] In June 2016, the US National Academies of Sciences, Engineering, and
Medicine released a report on their "Recommendations for Responsible Conduct" of gene drives.[25]
Models suggest that extinction-oriented gene drives will wipe out target species and that drives could
reach populations beyond the target given minimal connectivity between them.[26]
Kevin M. Esvelt stated that an open conversation was needed around the safety of gene drives: "In our
view, it is wise to assume that invasive and self-propagating gene drive systems are likely to spread to
every population of the target species throughout the world. Accordingly, they should only be built to
combat true plagues such as malaria, for which we have few adequate countermeasures and that offer
a realistic path towards an international agreement to deploy among all affected nations.".[27] He
moved to an open model for his own research on using gene drive to eradicate Lyme disease in
Nantucket and Martha's Vineyard.[28] Esvelt and colleagues suggested that CRISPR could be used to
save endangered wildlife from extinction. Esvelt later retracted his support for the idea, except for
extremely hazardous populations such as malaria-carrying mosquitoes and isolated islands that
would prevent the drive from spreading beyond the target area.[29]
History
Austin Burt, an evolutionary geneticist at Imperial College London, introduced the possibility of
conducting gene drives based on natural homing endonuclease selfish genetic elements in 2003.[6]
Researchers had already shown that such genes could act selfishly to spread rapidly over successive
generations. Burt suggested that gene drives might be used to prevent a mosquito population from
transmitting the malaria parasite or to crash a mosquito population. Gene drives based on homing
endonucleases have been demonstrated in the laboratory in transgenic populations of mosquitoes[30]
and fruit flies.[31][32] However, homing endonucleases are sequence-specific. Altering their specificity
to target other sequences of interest remains a major challenge.[3] The possible applications of gene
drive remained limited until the discovery of CRISPR and associated RNA-guided endonucleases such
as Cas9 and Cas12a.
In June 2014, the World Health Organization (WHO) Special Programme for Research and Training
in Tropical Diseases[33] issued guidelines[34] for evaluating genetically modified mosquitoes. In 2013
the European Food Safety Authority issued a protocol[35] for environmental assessments of all
genetically modified organisms.
Funding
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Target Malaria, a project funded by the Bill and Melinda Gates Foundation, invested $75 million in
gene drive technology. The foundation originally estimated the technology to be ready for field use by
2029 somewhere in Africa. However, in 2016 Gates changed this estimate to some time within the
following two years.[36] In December 2017, documents released under the Freedom of Information
Act showed that DARPA had invested $100 million in gene drive research.[37]
Control strategies
Scientists have designed multiple strategies to maintain control over gene drives.
In 2020 researchers reported the development of two active guide RNA-only elements that, according
to their study, may enable halting or deleting gene drives introduced into populations in the wild with
CRISPR-Cas9 gene editing. The paper's senior author cautions that the two neutralizing systems they
demonstrated in cage trials "should not be used with a false sense of security for field-implemented
gene drives".[38][39]
If elimination is not necessary, it may be desirable to intentionally preserve the target population at a
lower level by using a less severe gene drive technology. This works by maintaining the semi-defective
population indefinitely in the target area, thereby crowding out potential nearby, wild populations
that would otherwise move back in to fill a void.[40]
CRISPR
CRISPR[41] is a DNA editing method that makes genetic engineering faster, easier, and more
efficient.[42] The approach involves expressing an RNA-guided endonuclease such as Cas9 along with
guide RNAs directing it to a particular sequence to be edited. When the endonuclease cuts the target
sequence, the cell repairs the damage by replacing the original sequence with homologous DNA. By
introducing an additional template with appropriate homologues, an endonuclease can be used to
delete, add or modify genes in an unprecedentedly simple manner. As of 2014, it had been tested in
cells of 20 species, including humans.[5] In many of these species, the edits modified the organism's
germline, allowing them to be inherited.
In 2014 Esvelt and coworkers first suggested that CRISPR/Cas9 might be used to build endonuclease
gene drives.[5] In 2015 researchers published successful engineering of CRISPR-based gene drives in
Saccharomyces[43], Drosophila,[44] and mosquitoes.[45][46] All four studies demonstrated efficient
inheritance distortion over successive generations, with one study demonstrating the spread of a gene
drive into laboratory populations.[46] Drive-resistant alleles were expected to arise for each of the
described gene drives, however this could be delayed or prevented by targeting highly conserved sites
at which resistance is expected to have a severe fitness cost.
Because of CRISPR's targeting flexibility, gene drives could theoretically be used to engineer almost
any trait. Unlike previous designs, they could be tailored to block the evolution of drive resistance in
the target population by targeting multiple sequences within appropriate genes. CRISPR could permit
a variety of gene drive architectures intended to control rather than crash populations.
Applications
Gene drives have two main classes of application, which have implications of different significance:
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introduce a genetic modification in laboratory populations; once a strain or a line carrying the gene
drive has been produced, the drive can be passed to any other line by mating. Here the gene
drive is used to achieve much more easily a task that could be accomplished with other
techniques.
introduce a genetic modification in wild populations. Gene drives constitute a major development
that makes possible previously unattainable changes.
Because of their unprecedented potential risk, safeguard mechanisms have been proposed and
tested.[43][47]
Disease vector species
One possible application is to genetically modify mosquitoes, mice, and other disease vectors so that
they cannot transmit diseases such as malaria and dengue fever in the case of mosquitoes and tickborne disease in the case of mice.[48] Researchers have claimed that by applying the technique to 1%
of the wild population of mosquitoes, that they could eradicate malaria within a year.[49]
Invasive species control
A gene drive could be used to eliminate invasive species and has, for example, been proposed as a way
to eliminate invasive species in New Zealand.[50] Gene drives for biodiversity conservation purposes
are being explored as part of The Genetic Biocontrol of Invasive Rodents (GBIRd) program because
they offer the potential for reduced risk to non-target species and reduced costs when compared to
traditional invasive species removal techniques. Given the risks of such an approach described below,
the GBIRd partnership is committed to a deliberate, step-wise process that will only proceed with
public alignment, as recommended by the world's leading gene drive researchers from the Australian
and US National Academy of Sciences and many others.[51] A wider Outreach Network for Gene Drive
Research exists to raise awareness of the value of gene drive research for the public good.[52]
Some scientists are concerned about the technique, fearing it could spread and wipe out species in
native habitats.[53] The gene could mutate, potentially causing unforeseen problems (as could any
gene).[54] Many non-native species can hybridize with native species, such that a gene drive afflicting
a non-native plant or animal that hybridizes with a native species could doom the native species.
Many non-native species have naturalized into their new environment so well that crops and/or
native species have adapted to depend on them.[55]
Predator Free 2050
The Predator Free 2050 project is a New Zealand government program to eliminate eight invasive
mammalian predator species (including rats, short-tailed weasels, and possums) from the country by
2050.[56][57] The projects was first announced in 2016 by New Zealand's prime minister John Key and
in January 2017 it was announced that gene drives would be used in the effort.[57] In 2017 one group
in Australia and another in Texas released preliminary research into creating 'daughterless mice',
using gene drives in mammals.[58]
California
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In 2017 scientists at the University of California, Riverside developed a gene drive to attack the
invasive spotted-wing drosophila, a type of fruit fly native to Asia that is costing California's cherry
farms $700 million per year because of its tail's razor-edged ovipositor that destroys unblemished
fruit. The primary alternative control strategy involves the use of insecticides called pyrethroids that
kills almost all insects that it contacts.[19]
Wild animal welfare
The transhumanist philosopher David Pearce has advocated for using CRISPR-based gene drives to
reduce the suffering of wild animals.[59] Kevin M. Esvelt, an American biologist who has helped
develop gene drive technology, has argued that there is a moral case for the elimination of the New
World screwworm through such technologies because of the immense suffering that infested wild
animals experience when they are eaten alive.[60]
See also
Biological machines
Cas9
Cas12a
Meiotic drive
Genome editing
Population control
Sterile insect technique
Synthetic biology
Target Malaria
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Further reading
Esvelt KM, Gemmell NJ (November 2017). "Conservation demands safe gene drive" (https://www.
ncbi.nlm.nih.gov/pmc/articles/PMC5689824). PLOS Biology. 15 (11): e2003850.
doi:10.1371/journal.pbio.2003850 (https://doi.org/10.1371%2Fjournal.pbio.2003850).
PMC 5689824 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5689824). PMID 29145398 (https://
pubmed.ncbi.nlm.nih.gov/29145398).
Noble C, Adlam B, Church GM, Esvelt KM, Nowak MA (June 2018). "Current CRISPR gene drive
systems are likely to be highly invasive in wild populations" (https://www.ncbi.nlm.nih.gov/pmc/arti
cles/PMC6014726). eLife. 7: 219022. bioRxiv 10.1101/219022 (https://doi.org/10.1101%2F21902
2). doi:10.7554/eLife.33423.002 (https://doi.org/10.7554%2FeLife.33423.002). PMC 6014726 (htt
ps://www.ncbi.nlm.nih.gov/pmc/articles/PMC6014726). PMID 29916367 (https://pubmed.ncbi.nlm.
nih.gov/29916367). S2CID 196680955 (https://api.semanticscholar.org/CorpusID:196680955).
De Chant T (July 17, 2014). "Genetically Engineering Almost Anything" (https://www.pbs.org/wgb
h/nova/next/evolution/crispr-gene-drives/). NOVA. Retrieved 11 August 2014.
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Gene drive - Wikipedia
Johnson C (July 17, 2014). "Harvard scientists want gene-manipulation debate" (https://www.bost
onglobe.com/news/science/2014/07/17/harvard-scientists-propose-gene-technology-that-could-alt
er-organisms-wild/Ae4XBtXhwOLOeKPQlabbcP/story.html). National Geographic. Retrieved
11 August 2014.
Langin K (July 17, 2014). "Genetic Engineering to the Rescue Against Invasive Species?" (http://n
ews.nationalgeographic.com/news/2014/07/140717-gene-drives-invasive-species-insects-disease
-science-environment/). National Geographic. Retrieved 11 August 2014.
Zimmer C (July 17, 2014). "A Call to Fight Malaria One Mosquito at a Time by Altering DNA" (http
s://www.nytimes.com/2014/07/17/science/a-call-to-fight-malaria-one-mosquito-at-a-time-by-alterin
g-dna.html). The New York Times. Retrieved 20 July 2014.
"The age of the red pen" (https://www.economist.com/news/briefing/21661799-it-now-easy-edit-ge
nomes-plants-animals-and-humans-age-red-pen). The Economist. August 22, 2015. ISSN 00130613 (https://www.worldcat.org/issn/0013-0613). Retrieved 2015-08-25.
"The most selfish genes" (https://www.economist.com/news/briefing/21661801-giving-bits-dna-po
wer-edit-themselves-intriguing-and-worrying-possibility). The Economist. August 22, 2015.
ISSN 0013-0613 (https://www.worldcat.org/issn/0013-0613). Retrieved 2015-08-25.
Esvelt K. "Gene Drives for the Alteration of Wild Populations" (http://www.sculptingevolution.org/g
ene-drives/). Retrieved 11 August 2014.
External links
The Outreach Network for Gene Drive Research website (https://genedrivenetwork.org)
The Genetic Biocontrol of Invasive Rodents (GBIRd) program website (http://www.geneticbiocontr
ol.org/)
"Gene Drive from Harvard's Wyss Institute" (https://www.youtube.com/watch?v=Cy69C6vnFCQ/).
Wyss Institute. 2014-07-17. Retrieved 2014-08-11.
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