Content uploaded by Pedro Fidelman
Author content
All content in this area was uploaded by Pedro Fidelman on May 25, 2022
Content may be subject to copyright.
Environmental Science and Policy 135 (2022) 36–45
Available online 4 May 2022
1462-9011/© 2022 The Author(s). Published by Elsevier Ltd. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).
The principles driving gene drives for conservation
Sarah Hartley
a
,
*
, Riley Taitingfong
b
, Pedro Fidelman
c
a
University of Exeter Business School, Streatham Court, Exeter EX4 4PU, United Kingdom,
b
University of California San Diego, Herbert Wertheim School of Public Health and Human Longevity Science, 9500 Gilman Dr., La Jolla, CA 92093, USA
c
Centre for Policy Futures, The University of Queensland, Brisbane, QLD 4072, Australia
ARTICLE INFO
Keywords:
Conservation
Invasive species
Gene drive
Governance
Principles
Emerging technology
ABSTRACT
Gene drive technology is an emerging biotechnology with the potential to address some of the most intractable
global biodiversity conservation issues. Scientists are exploring potential gene drive applications for managing
invasive species and building resilience in keystone species threatened by climate change. The possibility to use
gene drive for these conservation purposes has triggered signicant interest in how to govern its development
and eventual applications. This includes a plethora of documents prescribing governance principles, which can
be a sensible response to the governance gap created by emerging technologies and help shore up legitimacy. We
conducted qualitative documentary analysis to examine the range and substance of principles emerging in the
governance of conservation gene drive. Such analysis aimed to better understand the aspirations guiding these
applications and how scientists and other experts imagine their responsibility in this eld. We found a collection
of recommendations and prescriptions that could be organised into a set of seven emerging principles intended to
shape the governance of gene drive in conservation: broad and empowered engagement; public acceptance;
decision-making informed by broad ranging considerations, state and international collaboration; ethical
frameworks; diverse expertise; and responsible self-regulation by developers. We lay bare these emergent
principles, analyzing the way in which they are valued, prioritized, and their strengths and weaknesses. By
identifying these prescriptive principles, stakeholders can further interrogate their merits and shortcomings and
identify more concrete ways that governance frameworks might embody them.
1. Introduction
The conservation of Earth’s biodiversity represents an accelerating
and intractable global issue. Despite the growth of conservation efforts
worldwide, biodiversity continues to decline at unprecedented rates and
across ecological scales (IPBES, Brondizio, 2019; Rands et al., 2010).
The monetary costs of invasive species worldwide was estimated to
reach US$162.7 billion in 2017 (Diagne et al., 2021).
Recent advancements in genome editing have enabled new possi-
bilities in the genetic modication of wild organisms, including for
purposes of conservation (Piaggio et al., 2017). One highly anticipated
advancement is gene drive, an emerging technique of genetic engineering
designed to rapidly spread traits through populations of sexually
reproducing organisms (NASEM, 2016). Using tools like CRISPR/Cas9,
certain gene drive systems can circumvent typical rules of genetic in-
heritance, facilitating the spread of genetic modications through nearly
100% of a population of organisms (Esvelt and Gemmell, 2017). In this
way, entire populations of organisms could be altered more rapidly and
effectively than ever before. Given the enormous challenge biodiversity
loss presents, scientists have started to explore the possibility of using
gene drive to solve conservation problems, particularly concerning
invasive species that contribute to biodiversity loss, or resilience in
keystone species threatened by climate change or other anthropogenic
pressures (Edwards et al., 2017).
Gene drive applications for conservation are appealing as they are
designed to spread a particular trait through an entire population, but
this design feature is also the source of signicant concern (Esvelt and
Gemmell, 2017). These concerns led to calls for gene drive governance
as early as 2014 (Oye et al., 2014). Certain governments and organiza-
tions are calling for moratoria on gene drives, and a wide variety of
commenters acknowledge that the technical development of gene drives
currently outpaces the capacity of existing mechanisms to govern their
use (Barnhill-Dilling et al., 2019; Kelsey et al., 2020; Rabitz, 2019). A
number of organizational bodies have engaged in efforts to develop
frameworks for gene drive governance (e.g., Convention on Biological
Diversity, 2017; NASEM, 2016; NIH, 2021; Redford et al., 2019).
* Corresponding author.
E-mail addresses: sarah.hartley@exeter.ac.uk (S. Hartley), rtaiting@ucsd.edu (R. Taitingfong), p.delman@uq.edu.au (P. Fidelman).
Contents lists available at ScienceDirect
Environmental Science and Policy
journal homepage: www.elsevier.com/locate/envsci
https://doi.org/10.1016/j.envsci.2022.04.021
Received 3 February 2022; Received in revised form 26 April 2022; Accepted 28 April 2022
Environmental Science and Policy 135 (2022) 36–45
37
However, no single entity is leading this effort and extant frameworks,
such as the Convention on Biological Diversity (CBD) and its Cartagena
Protocol may fall short in providing adequate oversight of gene drive
organisms, which pose unique issues with regards to risk assessment and
transboundary movement (Rabitz, 2019).
Gene drive technology presents a formidable challenge for the
governance of biotechnology (Sustainability Council of New Zealand,
2018). One of the dominant governance challenges is that gene drive
organisms are designed to spread through and possibly eliminate whole
populations or species. This means gene drive is a global, transboundary
technology that will not respect political or geographical boundaries and
will require international cooperation. The international response to this
challenge has started to take shape at the United Nation’s CBD and the
International Union for Conservation of Nature (IUCN) (Thizy et al.,
2020). The CBD offers a potential governance framework and its focus
on biodiversity conservation makes it a key actor for gene drive appli-
cations in conservation as gene drive is both designed as a solution to the
global biodiversity loss, and yet also presents a risk to biodiversity
(Reynolds, 2020). However, at both the IUCN and the CBD, gene drive
technology has provoked considerable controversy and become a deeply
divisive issue which may pose a signicant problem for international
gene drive governance (Reynolds, 2020).
Calls for engagement and consideration of ethical concerns are
familiar accompaniments to emerging technologies in lieu of interna-
tional or state laws and regulations. In this challenging international
environment, governance principles can be a sensible response to the
governance gap. They might also respond to a perceived democratic
decit in decision-making and as a means to shore up legitimacy (Ansell
and Torng, 2016). Indeed, gene drive has been accompanied by strong
discourses of responsibility (Stelmach et al., 2022), co-production
(Hartley et al., 2021a; Ledingham and Hartley, 2021) and public,
stakeholder and community engagement (Barnhill-Dilling and Del-
borne, 2021; NASEM, 2016). Gene drive has also triggered a plethora of
commitments, proclamations, recommendations and principles made by
gene drive developers, supporters and others (see Akbari et al., 2015;
Benedict et al., 2008; Benedict et al., 2018; Emerson et al., 2017; Long
et al., 2020; Min et al., 2018; Oye et al., 2014). Yet all of these refer to
gene drive more generally and are predominantly led by those devel-
oping or funding gene drive applications for global health.
Governance is a popular yet nebulous concept which reects the
growing trend of decentralized power (Ansell and Torng, 2016). In the
governance of emerging technologies such as gene drives, there is
frequently a ‘governance gap’ that emerges in the wake of the technol-
ogy (Marchant and Tournas, 2019). Here, policy-makers and regulators
must keep pace with the technology at a time when there is considerable
uncertainty, wrestle with the degree to which it requires new legislation
or regulations, and frequently have to co-ordinate with agencies in other
countries (Marchant and Tournas, 2019; Rabitz, 2019; Weiss Evans and
Palmer, 2018). Often, experts in the relevant eld ll this gap with a
proposal for a set of principles to guide governance, perhaps emerging
from a single meeting, organization or set of researchers. Examples
include the Organisation for Economic Co-operation and Development
Principles for Articial Intelligence (OECD, 2019), the Oxford Principles
for Geoengineering (Rayner et al., 2013), and International Principles
for Neurotechnologies (Marchant and Tournas, 2019). Such principles
can be an effective tool for ethical governance and have a key role to
play in the governance of emerging technologies (Littler, 2021).
To date, there has been no such set of explicit principles to ll the
governance gap for conservation gene drive. Instead, there has been a
urry of different prescriptions for responsible governance from expert
stakeholders and academics. Many of these focus on risk governance and
on gene drive mosquitoes designed to tackle global health challenges
like malaria, which experts expect will be the rst application of gene
drive tested in the wild (Esvelt and Gemmell, 2017). In addition, some
academics and organizations have proposed ethical principles for gene
drive more broadly (Annas et al., 2021; Emerson et al., 2017; Rudenko
et al., 2018). At this time, prescriptions for the governance of gene drive
for conservation in this mix remain unclear. These prescriptions need
disentangling, collating and examining in order to understand the
governance principles emerging in this space, who is leading such
governance efforts, and what might be missing. Further, as these prin-
ciples have the potential to shape the future (Rayner et al., 2013), it is
important to take stock of governance prescriptions for conservation
gene drive at a time when governance frameworks are still malleable.
Through a literature and documentary analysis, we propose to
address this gap in two ways. First, we identify the gene drive applica-
tions under development for conservation as well as those proposed for
future exploration to understand the range of gene drive applications in
need of governance. Second, we identify and analyze the recommen-
dations and principles that experts prescribe with the explicit aim to
shape the governance of conservation gene drive. We nd seven emer-
gent principles guiding gene drive research and development, which are
concerned with engagement, public acceptance, the breadth of gover-
nance, international collaboration, ethical input, the breadth of exper-
tise and self-regulation. We lay bare these emergent principles,
analyzing the way in which they are valued, prioritized, and their
strengths and weaknesses. Lastly, we explore the role of principles as a
site of governance. By revealing the ways in which expert stakeholders
prescribe and prioritize governance principles, we contribute to a more
open discussion of governance efforts in this space.
2. Gene drive applications in conservation
Various gene drive technologies are being developed and proposed
for biodiversity conservation. At the time of writing, just two key in-
terventions are in development: (1) sex-biasing gene drives to eradicate
invasive rodents from islands, and (2) underdominance systems to
suppress populations of mosquitoes that vector avian malaria to en-
dangered native birds in Hawai’i. Invasive rats and mice threaten island
ecosystems by way of predation and habitat disruption. Gene drives are
being explored as an alternative to the broad-spectrum toxicants
currently used for their eradication. Drive systems proposed for rodent
eradication include the non-transgenic t-Sry systems that produce
“daughterless mice”, which are only able to have male offspring.
CRISPR/Cas9 systems are still in development in rodents (Callaway,
2018; Leitschuh et al., 2018). The second intervention involves the Culex
quinquefasciatus, commonly known as the southern house mosquito,
vectors avian malaria (Plasmodium relictum) to Hawai’i’s endemic
honeycreepers. Warming temperatures are expanding the range of culex
into the high elevations where honeycreepers have previously escaped
infection, meaning transmission will intensify at lower elevations and
expand into higher elevations. Researchers at the University of Hawai’i
at M¯
anoa and Hilo are developing localized gene drives known as
under-dominance systems that could suppress mosquito populations by
engineering mosquitoes that are either unable to reproduce or transmit
the malaria parasite (Goldman, 2016).
At least eight other proposed interventions remain speculative or in
early stages of technical development. The rst of these involve coral
reefs which are threatened by a number of factors, particularly, the
impacts of a warming ocean as a result of anthropogenic climate change
(Hughes et al., 2017). Research is underway in Australia, Saudi Arabia,
and the UK to identify genes that control coral resistance to heat stress,
including the genetic engineering of coral bacteria, though these stra-
tegies remain speculative at this time (Cornwall, 2019). The second
involves grey squirrels (Sciurus carolinensis) which are invasive in the
UK, and contribute to the decline of native red squirrels (Sciurus vul-
garis) and impact woodlands by stripping bark from trees. With support
from the non-governmental organization, the European Squirrel Initia-
tive, scientists at the Roslin Institute have modelled self-limiting drive
systems known as HD-ClvR (a combination of homing, daisyeld, and
cleave-and-rescue gene drives) that could suppress squirrel populations
via a number of mechanisms including infertility or embryonic lethality
S. Hartley et al.
Environmental Science and Policy 135 (2022) 36–45
38
(Faber et al., 2021). The third speculative intervention involves the
crown-of-thorns starsh (Acanthaster spp.) which is a large coral-eating
starsh with unpredictable outbreaks in its natural range across the
Indo-Pacic region. Crown-of-thorns outbreaks are considered a leading
cause of coral reef decline, and are exacerbated by climate change as
their predation increases in water with lower pH. Genetic methods of
biocontrol are being considered to control crown-of-thorns starsh,
including CRISPR/Cas9 gene drives systems that might target repro-
ductive rates to suppress populations (Dumas et al., 2020; Høj et al.,
2018; Kamya et al., 2017; Redford et al., 2019).
Two of the remaining speculative interventions involve feral cats and
cane toads, both invasive species in Australia. Feral cats spread disease
and prey on endangered species all over the world, but in Australia,
where the feral cat population is estimated somewhere between 1.5 and
5.5 million, scientists are considering genetic biocontrol methods
including the use of sex-biasing gene drives to eradicate their pop-
ulations. However, more research is needed to develop successful
mammalian gene drives and gain necessary genetic and ecological
knowledge about feral cats in Australia (Kachel, 2018; Trouwborst et al.,
2020). Also in Australia, cane toads impact native ora and fauna by
poisoning organisms that eat them, and reducing prey populations for
native insectivores. Scientists are exploring w-shredder gene drives as a
mechanism to eradicate cane toads by sterilizing females or causing
them to produce only male offspring (Holman, 2019).
Scientists at the Roslin Institute at the University of Edinburgh are
exploring an infertility-carrying gene drive as a tool to manage invasive
signal craysh (Rudgard, 2021). First introduced to the UK from the US
in the 1970 s, signal craysh pose signicant threat to Britain’s native
white-clawed craysh by destabilizing their habitats with extensive
burrowing, and transmitting craysh plague, a disease with a 100%
mortality rate when contracted in the native craysh. Finally, scientists
in New Zealand are considering whether gene drives reducing fertility
could be a potential tool to control populations of invasive brushtail
possums and wasps. The brushtail possum is a marsupial introduced
from Australia that impacts New Zealand’s native plants, birds, and
invertebrates, and vectors bovine tuberculosis (Allot, 2021; Royal So-
ciety Te Ap¯
arangi, 2017). Though research to understand possum ge-
netics is underway, little is known about functional genetics in
marsupials, and a gene drive could prove challenging as it would require
that very large amounts of gene drive-carrying possums (between 1%
and 10% of the wild population) be bred and released. Scientists at the
University of Wellington and University of Otago are exploring a gene
drive targeting spermatogenesis to control the common wasp (Vespula
vulgaris) (Lester et al., 2020). The common wasp is one of two species of
invasive wasps in New Zealand (the other is Vespula germanica, the
German Wasp) that pose conservation challenges through predation on
native insects, and feeding on beech forest honeydew which is an
important food source for native birds, bats, insects, and lizards.
Containment is of particular concern in relation to both possum and
wasp applications in New Zealand as the brushtail possum is a protected
species in the neighbouring Australia, and Vespula wasps are an
important part of ecosystems throughout Europe.
3. Methods
We used document analysis, an inductive and exploratory qualitative
approach that aims to “access a particular perspective in depth, rather
than to test a specic hypothesis” (Sovacool et al., 2018). Document
analysis has been widely pursued in the context of emergent healthcare
policies, as documents can provide material for study prior to or in the
absence of material changes in or adoption of new policies (Shaw et al.,
2006). Given the nascent state of conservation gene drive, governance
documents, such as the ones we analyse here, provide a valuable way to
investigate emerging principles that may eventually inform the devel-
opment of policies and legislation.
We began our identication of the data set in December 2020 with
broad search of literature addressing conservation gene drive technol-
ogies published up to January 2021. Given our interest in emerging
governance principles, we identied two criteria for inclusion in our
analysis. All included documents were to address: (1) conservation gene
drives, and (2) some topic related to the governance of conservation
gene drive technologies. Our initial searches returned 41 documents. We
eliminated documents that did not satisfy one or both of our criteria.
Many were eliminated because they discussed topics related to the use of
genetic modication or synthetic biology for environmental conserva-
tion, but did not address gene drives specically (n =14). For example,
(Redford et al., 2014) discuss the potential application of synthetic
biology to biodiversity conservation, but do not discuss gene drives.
Others were eliminated because conservation gene drives were only
briey mentioned within a broader discussion of gene drives (n =12).
For example, Thizy et al. (2020) review the regulatory landscape around
gene drive but do not focus exclusively on conservation. One was
eliminated because it featured predominantly technical discussion of a
conservation gene drive for wasps but had no mention of governance
(Lester et al., 2020), and one because it did not address particular ap-
plications of gene drive technologies (Golnar et al., 2020). This left us
with 13 documents that focused more narrowly on conservation gene
drives, and included some mention of topics related to their governance
(e.g., risk assessment, engagement, regulatory processes). Lastly, we
identied 5 additional eligible articles through a nal database search
and by reviewing references in the included literature.Table 1.
Data analysis was an iterative and reexive process of thematic
analysis conducted in early 2021 (Braun and Clarke, 2019). Collabora-
tively, we established a process through which to code systematically
the key governance principles communicated in the dataset and itera-
tively generate themes. Following Braun and Clarke we emphasize that
themes were reexively generated rather than pre-existing naturally
within the data (Braun and Clarke p. 593). After nalizing the dataset,
all three authors independently coded two documents. This involved a
three-step process: (1) close reading of each document to locate dis-
cussion on governance, (2) pulling quotations from relevant sections to
capture the governance principles communicated in each document, and
(3) distilling those quotations into “codes” that succinctly describe the
governance principle (e.g., “community engagement,” “deliberation,”
“public acceptance”). [Name removed for review] then coded the whole
data set between December 2020 and January 2021 guided by monthly
coding meetings held remotely across three countries (USA, UK and
Australia). Upon completion of coding, we had generated more than 40
codes that captured both recurrent and unique governance principles
communicated across the documents. [name removed for review] and
[name removed for review] reviewed the codes, then [name removed for
review] pr´
ecised the codes into nine higher level themes. The authors
then met to discuss the themes. After identifying points of overlap across
some of the initial themes, the authors further narrowed them down to
seven.
We understand themes as “patterns of shared meaning” which were
underpinned by the core concept of governance principles (Braun and
Clarke, 2019, p. 593). In the interest of capturing broad themes in the
discussion on governance, we conceived of governance principles as
encompassing both descriptive comments and prescriptive guidelines on
conservation gene drive governance. That is, we coded selections mak-
ing explicit calls for particular approaches to governance as well as se-
lections observing or exploring topics of conservation gene drive
governance that did not necessarily advocate a particular approach. For
instance, while some articles described international policies and regu-
lations that may be relevant for the governance of conservation gene
drive organisms, others made more explicit arguments for particular
practices or approaches to governance, and others yet involved a com-
bination of both.
S. Hartley et al.
Environmental Science and Policy 135 (2022) 36–45
39
4. Seven governance principles in conservation gene drive
Here, we lay out the seven governance principles identied in the
data according to prevalence (i.e., the frequency with which they
occurred in the analyzed documents) (Table 2). For principles to ll the
‘governance gap’, they must be both ambitious and capable of practical
application (Cohen-Shacham et al., 2019). Therefore, we analyze their
strengths and weaknesses and discuss their aspirations and potential for
practitioners.
4.1. Governance must involve inclusive, broad and empowered
engagement
This principle was the most widespread, with thirteen of eighteen
documents addressing the need for conservation gene drive governance
to involve public or community engagement
1–8, 11–13, 15, 18
. The docu-
ments commonly call for forms of engagement that go beyond unidi-
rectional activities like education or public comment, with a few
explicitly addressing the limits of such activities
4, 8
. One of these chal-
lenged the assumption that more education leads to more approval,
stating that “managers often attribute opposition [to gene drives] to
ignorance, but studies show that public education designed specically
to garner support does not necessarily increase acceptance and can
heighten conict”
8
. In light of this insight, authors of this article
asserted that “public engagement campaigns should be at least as much
about public deliberation as they are public education”
8
.
Similarly, an article on public opinion surrounding the use of gene
drive for pest control cautioned the use of “information-driven”
engagement, stating that “future public engagement regarding gene
drive that relies solely on educating people by providing more scientic
information and facts seriously risks limiting (and potentially inaming)
public discourse”
4
. This article called for “values-based” engagement
given ndings that “public attitudes towards gene drive are not formed
on scientic knowledge or demographics alone, but are heavily inu-
enced by underlying worldviews, which encapsulate a broad and
interactive system of attitudes, beliefs, and values.”
Overall, the discussion on engagement demonstrated broad calls for
deliberative forms of engagement, and emphasized the need to account
Table 1
Documents focused on gene drive in conservation and discussing governance.
No. Document Type Pages
1 Webber, B. L., Raghu, S., & Edwards, O. R. (2015)
Opinion: Is CRISPR-based gene drive a biocontrol
silver bullet or global conservation threat? PNAS,
112(34): 10565–10567.
Opinion
article
3
2 Edwards, O., Brown, P., Tizard, M., Strive, T., &
Sheppard, A. Taking a responsible approach to new
genetic technologies for conservation. CSIRO.
(2017). https://ecos.csiro.au/taking-responsible-
approach-new-genetic-technologies-conservation/
Web article 4
3 Esvelt KM, Gemmell NJ (2017) Conservation
demands safe gene drive. PLoS Biol 15(11):
e2003850. https://doi.org/10.1371/journal.
pbio.2003850
Article 8
4 Harvey-Samuel, T., K. J. Campbell, M. Edgington
et al. 2017. Trialling gene drives to control invasive
species: What, where and how? In Island Invasives:
Scaling Up to Meet the Challenge, Proceedings of the
International Conference on Island Invasives 2017,
C. R. Veitch, M. N. Clout, A. R. Martin, J. C. Russell,
and C. J. West, eds. Gland, Switzerland: IUCN
Species Survival Commission, 618–627.
Article 10
5 Piaggio, A. J., Segelbacher, G., Seddon, P. J., Alphey,
L., Bennett, E. L., Carlson, R. H., Friedman, R. M.,
et al. Is it Time for Synthetic Biodiversity
Conservation? (2017). Trends in Ecology &
Evolution, 32(2): 97–107.
Article 11
6 Dearden, PK, Gemmell, NJ, Mercier, OR, Lester, PJ,
Scott, JM, Newcomb, TRB, Jacobs, JME, Goldson,
SJE, & Penman, DR. (2018). The potential for the use
of gene drives for pest control in New Zealand: a
perspective, Journal of the Royal Society of New
Zealand, 48:4, 225–244, DOI: 10.1080/
03036758.2017.1385030
Article 21
7 McFarlane GR, Whitelaw CBA, Lillico SG. CRISPR-
based gene drives for pest control. Trends Biotechnol
2018;36(2):130–133.
Article 4
8 Moro, D., Byrne, M., Kennedy, M., Campbell, S and
Tizard, M. (2018). Identifying knowledge gaps for
gene drive research to control invasive animal
species: The next CRISPR step. Global Ecology 55
and Conservation, 13, e00363.
Article 15
9 Barnhill-Dilling, K., Serr, M., Blondel, V.D., Godwin,
J. Sustainability as a Framework for Considering
Gene Drive Mice for Invasive Rodent Eradication.
(2019). Sustainability, 11, 1334.
Article 12
10 Godwin, J., Serr, M., Barnhill-Dilling, S. K., Blondel,
D. V., Brown, P. R., Campbell, K., Delborne, J., 3
Lloyd, A. L., Oh, K. P., Prowse, T. A. A., Saah, R. and
Thomas, P. (2019). Rodent gene drives for
conservation: opportunities and data needs.
Proceedings of the Royal Society B: Biological
Sciences, 286, 20191606.
Article 9
11 Kohl, P. A., Brossard, D., Scheufele, D. A., & Xenos,
M. A. (2019). Public views about editing genes in
wildlife for conservation. Conservation Biology:
1–10. Doi:10.1111/cobi.13310
Article 10
12 Redford, K.H., Brooks, T.M., Macfarlane, N.B.W. and
Adams, J.S. (eds.) (2019). Genetic frontiers for
conservation: An assessment of synthetic biology and
biodiversity conservation. Technical assessment.
Gland, Switzerland: IUCN. xiv +166pp.
Report 184
13 Revive & Restore. Ocean Genomics Horizon Scan.
(2019).
Report 168
14 Testbiotech comment on the IUCN report “Genetic
frontiers for conservation, an assessment of synthetic
biology and biodiversity conservation” (2019).
Letter 11
15 MacDonald, E. A., Balanovic, J., Edwards, E. D.,
Abrahamse, W., Frame, B., Greenaway, A., …
Tompkins, D. M. (2020). Public Opinion Towards
Gene Drive as a Pest Control Approach for
Biodiversity Conservation and the Association of
Underlying Worldviews. Environmental
Communication, 1–15.
Article 16
16 Article 11
Table 1 (continued )
No. Document Type Pages
Palmer, S, Mercier, OR, & King-Hunt, A. (2020).
Towards rangatiratanga in pest management? M¯
aori
perspectives and frameworks on novel
biotechnologies in conservation. Pacic
Conservation Biology, CSIRO.
17 Reynolds, J. 2020. Governing New Biotechnologies
for Biodiversity Conservation: Gene Drives,
International Law, and Emerging Politics. Global
Environmental Politics, 20 (3). https://doi.org/
10.1162/glep_a_00567
Article 21
18 Serr, M. E., Valdez, R. X., Barnhill-Dilling, K.S.,
Godwin, J., Kuiken, T., Booker, M. (2020). Scenario
analysis on the use of rodenticides and sex-biasing
gene drives for the removal of invasive house mice
on islands. Biological Invasions, 22: 1235–1248.
Article 14
Table 2
Emergent principles guiding gene drive research and development for conser-
vation applications, in order of prevalence.
1. Governance must involve inclusive, broad and empowered engagement
2. Gene drive research and development must have public acceptance
3. A broad range of considerations must be incorporated in decision-making
4. States and international actors must collaborate and act
5. Governance must draw on broad frameworks which are attentive to ethical principles
6. A broad range of experts must provide input
7. Developers must self-regulate and respond responsibly
S. Hartley et al.
Environmental Science and Policy 135 (2022) 36–45
40
for diverse public or stakeholder values. For instance, authors called for
“open debates that engage diverse expert and lay voices”
3
, “balanced
deliberation that engages multiple stakeholders”
7
, “robust, informed,
and deliberative engagement” with stakeholders
12
, and consideration of
how deliberative engagement can “meaningfully inuence decision-
making"
15
. Two documents called for iterative approaches to engage-
ment
3, 7
such that ”affected societies” can “continually revisit issues,
[and] reframe their views in the light of subsequent experience”
3
, and
clearly communicate “social and ethical priorities” and a “full range of
standards that genetic interventions must meet to be considered for
environmental release”
7
. While most documents focused on broad public
engagement, or the engagement of “all” affected societies, a few focused
instead on specic stakeholder groups such as local communities and/or
Indigenous peoples
1, 6, 18
. Just one document took up this conversation
in depth, calling for the “meaningful engagement” of M¯
aori in Aotearoa
New Zealand, and specically discussing how "M¯
aori values and con-
ceptual frameworks [can] give M¯
aori more agency to engage with
contemporary issues like the role of biotechnologies in relation to
environmental threats"
18
.
4.2. Gene drive research and development must have public acceptance
Eleven documents address the role of public acceptance within
conservation gene drive governance
1, 2, 3, 6, 8, 11, 14, 15, 16–18
. This dis-
cussion is characterized by broad calls to cultivate communication,
trust, transparency, and openness across scientists, governments, and
communities or broader publics, though recommendations about how
best to achieve this vary. For instance, some authors acknowledge the
need for broad public acceptance prior to release of a gene driv-
e–modied organism for conservation
2, 14, 17, 18
, while others call more
generally for attention to issues of “social acceptance” or “community
acceptance,” without necessarily prescribing what that should entail
15,
16
.
Two contested concepts that emerged in this discussion were social
acceptance (also social license) and consent. Social license is described in
one document as “consent of communities” for various activities,
emerging originally in connection to commercial tourism and extractive
practices like mining, later getting adopted within agriculture, forestry,
and now biotechnology
18
. The documents problematize the concept of
social license as prioritizing “industry prot targets” over community
needs or aspirations, and potentially circumnavigating processes that
perhaps ought to be regulated by governments. Other calls to “socialize”
gene drive technologies emphasizing the role of information in
achieving social acceptance, particularly information surrounding the
risks, benets, and efcacy of conservation gene drives
16
, as well as
biological, genetic, and ecological information that would inform the
development of a task like risk assessment
17
.
Other documents focused more on the concept of consent
6, 15, 18
. The
IUCN report
6
commented broadly on stipulations of free, prior, and
informed consent under international legislation, namely the Conven-
tion on Biological Diversity and Cartagena Protocol. Such reports and
other documents
15, 18
express consent as necessary in the context of
conservation gene drives, while recognizing the challenges and limita-
tions of extant models. The IUCN report
6
acknowledges that some of the
most active states in biotechnology are not signatories to the Cartagena
Protocol, and Palmer et al.
18
note that both concepts – social license and
free, prior, and informed consent – operate in a Western paradigm, and
may exclude Indigenous worldviews.
4.3. A broad range of considerations must be incorporated in decision-
making
More than half of the documents expressed the need for conservation
gene drive governance to consider a broad range of factors in decision-
making
1, 2, 6, 7, 8, 9, 10, 11, 12, 18
. This discussion framed the complexity
1, 8,
9, 12
of both conservation issues and emerging gene drive technologies,
and called for decisions to attend to technical, social, economic, and
environmental considerations alike. Technological complexity was
described as arising from the interconnected issues of “large scale,
ecological, and systemic” risks
12
, “multiplex interrelations with the
closer and wider environment,” and challenges of achieving spatio-
temporal controllability and anticipating evolutionary effects
9
. Such
discussion underscored actions to ensure the safety and efcacy of con-
servation gene drives, namely through sound scientic risk assessment
and mitigation of biosecurity threats. Recommendations emphasized
continued scientic research to account for technical complexity. For
instance, one document stated that the only way to manage risks of
pursuing gene drive (as well as the risks of not pursuing them) is “to have
more information, more research and more evidence with which to
better understand and mitigate those risks and proceed responsibly”
2
.
Social complexity was acknowledged as the diverse social, economic,
and political contexts in which community stakeholders exist
1,8
, and
acknowledgment that those stakeholders have diverse perceptions of
what constitutes risk, and values regarding human relationships to na-
ture
1
. For example, an article discussing the use of gene drive mice for
invasive rodent eradication notes that gene drives “emerge in complex
human systems” and identies “the potential for socio-political oppo-
sition about the environmental release of gene drive rodents” as one of
the most signicant potential barriers to their use
1
. Just one document
provided expanded discussion of how to integrate the views and values
of a particular stakeholder group (Indigenous people in Aotearoa New
Zealand) into decision-making, taking time to describe specic M¯
aori
values and legal frameworks that may be relevant for the context of
conservation biotechnology including gene drives
18.
4.4. States and international actors must collaborate and act
Half of the documents identify the need for conservation gene drive
governance to involve state and international collaboration, involve-
ment, and action
1, 5–7, 12, 13, 14, 15, 18.
A major theme in this discussion is
the tension between local and national governance of gene drives. While
all articles recognized the potentially global ramications of gene drive
organisms, two positioned local community governance as central to
decision-making
1, 18
and others focused on the need for broader inter-
national governance given the possibility of transboundary movement
6,
12, 13, 15
. For instance, an article discussing the use of gene drive mice to
eradicate invasive rodents from islands asserted that “local community
governance should be central to related decisions” given that local
communities will bear “the most direct effects of gene drive organ-
isms”
1
. Another article on gene drive for invasive species management
focused more squarely on international consequences of self-
propagating drives, cautioning that such drives should not be released
unless “international spread is the explicit goal”. This piece questioned
the use of self-propagating gene drives for conservation altogether,
expressing the view that they “should only be built to combat true pla-
gues such as malaria, for which we have few adequate countermeasures
and there is a realistic path towards and international agreement to
deploy among all affected nations”
13
.
Other documents emphasizing the need for the international
collaboration called for the development of new forms of multilevel
governance that spans the local and global
6, 12, 15
. Two documents
offered an extensive discussion on state and international regulatory
systems relevant to conservation gene drive (e.g., CBD, Nagoya, Carta-
gena, Aarhus, UNDRIP, FPIC, ILO), with both agging intergovern-
mental coordination as a challenge for gene drive modied organisms
6,
12
. A minor but compelling theme in this discussion was the need to
develop protocols of liability and redress related to transboundary harm.
The IUCN report suggested that while “national and international legal
systems may provide for liability for environmental damage attributable
to synthetic biology”, including gene drives, there are “few international
frameworks that explicitly provide for liability – either on the part of
states or the part of operates – in the context of biosafety.” The report
S. Hartley et al.
Environmental Science and Policy 135 (2022) 36–45
41
references the Nagoya-Kuala Lumpur Supplementary Protocol on Lia-
bility and Redress (a binding supplementary agreement to the Cartagena
Protocol on Biosafety) as one example of state responsibility for inter-
national harm caused by living modied organisms
6
.
4.5. Governance must draw on broad frameworks which are attentive to
responsible and ethical principles
Eight of the eighteen documents identied the need for some kind of
broad framework to guide governance of conservation gene drives
1, 2, 5,
6, 9, 12, 15, 18
. The most recurrent theme in this discussion was re-
sponsibility, though descriptions of responsible governance varied. For
instance, two journal articles described responsible governance as that
which accounts for transboundary risks and seeks intergovernmental
collaboration. One article referenced the NASEM (2016) report on gene
drives to describe responsible governance as “international and inclu-
sive” governance that “anticipate[s] transboundary effects of gene-drive
modied organisms”
12
. The other article described “responsible and
ethical governance” as encompassing local and global regulation as well
as deliberative community engagement
15
. Other documents character-
ized responsibility as technological innovation for the public good
2
, with
some calling for limited adoption of precautionary approaches empha-
sized by the Cartagena Protocol and the CBD, for concern that precau-
tion may limit continued innovation
1,5
. For example, one opinion piece
in a scientic journal says that “responsible stewardship rejects positions
that forsake potential benets in deference to absolute caution and po-
sitions that ignore reasonably foreseeable risks to allow unfettered sci-
entic exploration”
5
.
A minor theme was sustainability. Two documents called for broad
governance frameworks that emphasize sustainability, with both
describing sustainability as governance that equally attends to the
environment, economy, and society to integrate complexity and uncer-
tainty
1, 6
. One of these documents was the IUCN report, which refer-
enced the World Commission on Environment and Development’s
concept of sustainable development as that which recognizes the inter-
dependence of economic and social development with environmental
conservation
6
. The IUCN report also dened sustainable development in
connection to principles of intergenerational and intragenerational eq-
uity, with the former “entail[ing] an obligation of stewardship of the
natural environment for future generations” and the latter emphasizing
“the need to meet the basic needs of current generations across cir-
cumstances and regions”
6
.
Three documents called for governance that is attentive to equity and
justice
1, 6, 18
. For instance, one journal article called for attention to
questions of fairness or justice, namely “who benets” from conserva-
tion gene drive technologies and “who gets to make decisions about
them"
1
. In addition to inter- and intra-generational equity, the IUCN
report also identied ”access to justice in environmental matters”, a
relevant principle for conservation gene drive governance
6
. It invokes
the Aarhus Convention to dene the principle of access to justice in
environmental matters, stating that: “any person – which includes any
environmental organisation – who considers their rights violated or in-
terests affected by an environmental decision has access to a court or
other independent and impartial review procedure to bear challenge the
substantive and procedural legality of the decision”
6
.
Finally, three documents discussed governance frameworks that ac-
count for the rights of Indigenous peoples
6, 12, 18
. The IUCN report and
one journal article discussed the free, prior, and informed consent
(‘consent’) of local and indigenous peoples, citing comments emerging
from the 2018 Convention on Biological Diversity’s Convention of
Parties that the “free, prior and informed consent of indigenous peoples
and local communities might be warranted when considering the
possible release of organisms containing engineered gene drives that
may impact their traditional knowledge, innovation, practices, liveli-
hood and use of land and water”
12
. One document problematized the
dominance of Western paradigms including consent in discussions on
governance of conservation biotechnology, cautioning that these may
exclude Indigenous worldviews
18
. Written from the context of Aotearoa
New Zealand, this article called for the adoption of M¯
aori legal frame-
works and customary protocols to inform the use of novel bio-
technologies in conservation.
4.6. A broad range of experts must provide input
Seven documents emphasized the need for broad expert input to
inform some aspect of decision-making surrounding conservation gene
drives
3, 5, 10, 11, 12, 15, 16
. Overall, this discussion advocated for increased
collaboration across technical experts in order to address knowledge
gaps and inform more risk assessment. For example, one article identi-
ed an “immediate need for the conservation community to more fully
engage with synthetic biology” in order to apply “their expertise to
robust risk assessments […], advise synthetic biologists of environ-
mental concerns and issues […], head off possibly ecologically
damaging initiatives […], and to identify the most appropriate conser-
vation problems for the development and implementation of acceptable
synthetic biology solutions.”
5
.
Another article called on “practitioners of CRISPR-Cas9 gene drive
[to] consider the lessons learned from decades of carefully regulated
CBC research”
10
, and yet another stated that appropriate decisions
require input from a broad range of “scientic disciplines” such as
“conservationists identifying potential targets, ecologists advising on the
biological appropriateness of these targets and efciencies of different
gene drive strategies, molecular biologists advising on the feasibility of
building proposed designs, mathematical modellers devising the most
efcient means of deploying these systems and, nally, managers who
will ultimately advise on the logistic feasibility of deployment.
11
.
An emergent theme in this discussion was interdisciplinarity, how-
ever, recommendations advocated exclusively for expanded collabora-
tion across scientic disciplines. This included calls for increased
communication between synthetic biologists and conservationists
5
; be-
tween conservationists, ecologists, molecular biologists, mathematical
modellers, and managers
11
; or for the integration of regulatory insights
and approaches to risk assessment from Classical Biological Control
10
.
4.7. Developers must self-regulate and respond responsibly
Finally, seven documents featured the nal theme on the need for
developers of conservation to respond through self-regulation and re-
sponsibility
2, 6, 7, 11, 13, 14, 16
. This discussion focused on the re-
sponsibility of gene drive developers to adopt a variety of genetic and
environmental strategies to limit the risk of indenite spread of gene
drive–modied organisms. Genetic strategies included the use of “low-
risk” drives (e.g., homing-based drives)
2
, self-limiting drives, “rigorous
and transparent safety testing in the laboratory”, and even complete
avoidance of the development of self-propagating drives
13
.
Recommended environmental controls included the development of
gene drive–modied organisms in laboratories far from existing pop-
ulations of the target species
13
, and the use of “small, isolated islands”
for the rst trials of gene drives, in concert with biosecurity measures to
restrict trafc to and from trial islands
11, 16
. The two documents that
recommended the use of islands for trials cited recommendations from
extant guidance reports (i.e., the World Health Organization’s 2014
guidance framework for testing of genetically modied mosquitoes and
the National Academies of Sciences, Engineering, and Medicine’s 2016
“Gene Drives on the Horizon” report) to emphasize that islands can
“maximise containment and efcacy”
11
and that “biosecurity risk can be
managed with some condence”
16
. One of these articles also identied
“social criteria” for eld trial site selection, including selection of “un-
inhabited areas which are not of great cultural value,” and the “presence
of a “credible regulatory structure and an enthusiastic local participant
(e.g. academic researcher or wildlife management agency) which
expertise regarding the invasive being targeted”
11
.
S. Hartley et al.
Environmental Science and Policy 135 (2022) 36–45
42
5. Discussion
Ansell and Torng (2016: 4) dene governance as “the interactive
processes through which society and the economy are steered towards
collectively negotiated objectives”. Governance is seen as distinct from
government where governing is achieved through traditional in-
stitutions (Rudenko et al., 2018). It is a looser collection of networks,
opening up opportunities for a more diverse set of actors to shape
policy-making and produce ‘desired outcomes’ for society. Governance
is important for emerging technologies, particularly in the assessment
and management of risk but increasingly in the shaping of technology
trajectories through approaches such as responsible innovation and
co-production (Hartley et al., 2019). Governance frameworks and
practices can exert signicant inuence on the types of science and
innovation brought into the world (Jasanoff, 2004). In recent decades,
there has been a widespread commitment to ‘opening up’ the gover-
nance of science and technology to enable a broader range of voices to
shape the developmental and design trajectory of emerging technologies
(Hagendijk and Irwin, 2006; Stirling, 2008). This opening up helps the
alignment of technological innovations with societal values (Kaebnick,
2021). In addition to this democratic or normative function, opening up
also increases epistemic diversity to increase the chance that an
emerging technology will be able to address a specic societal challenge
(Hartley et al., 2019; Stirling, 2010; Wynne, 1992).
Gene drive is an emerging technology that has been ‘opened up’
through explicit and public commitments to responsibility, engagement
and knowledge co-production (Hartley et al., 2021a; Ledingham and
Hartley, 2021). Specic principles and codes of ethics are starting to
emerge in order to guide gene drive developers, funders and supporters.
For example, Emerson et al. (2017) present a set of ve principles for
gene drive research with a focus on global health. They align their
principles with the recommendations of the NASEM report, arguing that
gene drive research must ‘promote the public good’, ‘promote stew-
ardship, safety, and good governance’, ‘demonstrate transparency and
accountability’, ‘engage thoughtfully with affected communities,
stakeholders, and publics’, and ‘foster opportunities to strengthen ca-
pacity and education’. More recently, Annas et al. (2021) offer a code of
ethics intended for adoption by gene drive scientists committed to
ethical guidance of gene drive research. Their code is organized around
three key values: ‘scientic responsibility’ (i.e., safety, security, and peer
review), ‘ecological stewardship’ (risk assessment, sustainability, and
conservation of biodiversity), and ‘public engagement and benet
sharing’ (loosely dened as transparency, fair distribution of risks and
benets, and informed agreement of impacted populations or commu-
nities). These efforts have been supported by other actors, particularly
the Foundation for the National Institutes of Health in the form of closed
forums, open webinars and panel discussions on gene drive governance
and particularly on the ethical principles guiding gene drive research (e.
g. Littler, 2021). However, governance principles can also be developed
as a form of self-governance designed to quell public concerns about an
emergent technology. For example, Gardiner and Fragni`
ere (2018)
critique the Oxford Principles for Geoengineering as being too instru-
mental and Ulnicane et al. (2020) highlight the criticism levelled at
experts’ attempts to address articial intelligence’s inherent contro-
versies with ethical principles, particularly perceived conicts of inter-
est and instrumental motivations to avoid codied regulations. Public
concerns about such conicts and motivations may undermine princi-
ples and increase concern about the technology.
We set out to explore emergent principles for gene drive in conser-
vation or invasive species management in order to understand whether a
distinct set of principles is emerging in this space. We examined the
range and substance of principles emerging in the development, testing
and governance of gene drive for conservation to better understand the
aspirations guiding these applications and how scientists and other ex-
perts imagine their responsibility in this eld. We found a collection of
recommendations and prescriptions related to the governance of gene
drive for conservation that could be organised into a set of seven themes.
These themes constitute a loosely dened set of emerging principles
intended to shape the governance of gene drive in conservation.
Many, if not all, sets of principles for emerging technologies call for
engagement. Such calls characterise the eld of gene drive with a sig-
nicant number of developers, funders and regulators committing to
public, stakeholder and/or community engagement. Similar to the
existing principles developed by Emerson et al. (2017) and Annas et al.
(2021) which focus of global health applications, in our case, engage-
ment is an important principle. In fact, engagement dominates the rst
three principles we identied, calling for governance to be inclusive, for
research to secure public acceptance, and for a broad range of consid-
erations to be incorporated in decision-making. However, despite this
dominant theme, there is considerable lack of clarity on the matter of
engagement. It is not clear who should be engaged, when and how they
should be engaged, and for which purpose. Although NASEM (2016)
makes clear engagement might include communities (e.g. people who
live close to a eld trial site), stakeholders (e.g. people with an interest
or a stake in gene drive), and lastly, publics (e.g. people indirectly
affected or interested in gene drive), those prescribing engagement in
gene drive often entangle these different groups and fail to specify which
groups need to be engaged, when or how (Hartley et al., 2022).
The documents are clear that engagement should go beyond
educating and informing people and should include deliberative forms
of engagement, yet the question of empowerment is rarely addressed.
For more than half a century, Arnstein’s (1969) ladder of citizen
participation has been a tool for evaluating the degree to which those
being engaged are empowered to shape decision-making. In the broader
debate on gene editing, Burall (2018: 438) has observed there has been
‘little attempt to engage the public on the implications of the technology
in a way that could alter the decisions of scientists and policymakers’.
However, the analysed documents have little to say on empowerment
and the degree to which people should have the power to decide on the
type of gene drive developed for conservation purposes, or whether a
eld trial should be approved by publics or local communities or not.
Indeed, the documents reveal an unhelpful division between “expert”
and “lay” or “public” stakeholders. For instance, one document calls for
open debate among “diverse expert and lay voices” (3) and another for a
panel of “diverse experts and stakeholders” (12). It is worth considering
how rigid designations of scientists as expert and non-scientists as “lay”
may foreclose consideration of expertise that exists in non-scientic or
non-technical communities and the degree to which they might be
empowered in decision-making.
Gene drive in conservation will bring a new set of actors to the
governance table, particularly actors who may not have been as relevant
in the governance of global health applications of gene drive. One set of
such actors are Indigenous peoples who may have legal responsibility for
land management and be deeply involved and invested in conservation
and invasive species management. Given the CBD’s instruction that gene
drives should be developed in consultation with Indigenous peoples and
the existing role of these peoples in land management and conservation,
we expected greater attention to the role of Indigenous actors than might
exist in principles for gene drive in global health. However, Indigenous
peoples are rarely addressed explicitly in the documents. Indeed, this is a
relatively homogenous group of documents with only one offering any
specicity about how or why Indigenous actors might be engaged
(Palmer et al., 2020). Further, there is very little discussion of the his-
toric and ongoing power imbalances that exist between Indigenous
peoples and those developing governance frameworks, such as govern-
ments, regulators, funders and developers.
Addressing engagement of Indigenous peoples and Indigenous rights
to self-determination would add to a growing conversation on Indige-
nous participation in deliberation and decision-making processes sur-
rounding gene drives (George et al., 2019; Hudson et al., 2019; NASEM,
2016; Taitingfong, 2019) and genetic engineering technologies more
broadly (Barnhill-Dilling and Delborne, 2019; Barnhill-Dilling et al.,
S. Hartley et al.
Environmental Science and Policy 135 (2022) 36–45
43
2020; Taitingfong and Ullah, 2021). Discussions on conservation gene
drive governance stand to benet from deeper engagement of this
literature, as it addresses contested concepts, such as consent and
engagement, which emerged in the conservation gene drive documents.
For instance, Hudson et al. (2019), Barnhill-Dilling and Delborne
(2019), and Taitingfong and Ullah (2021) discuss and model approaches
to Indigenous community engagement in the context of genetic engi-
neering. Lastly, scholars ought also to look to robust bodies of scholar-
ship written by Indigenous scientists and scholars generating critical
analyses of coloniality as it structures the development and governance
of other genetic and genomic technologies (Arvin, 2019; Claw et al.,
2018; Fox et al., 2020; TallBear, 2013; Tsosie and Claw, 2019).
Another group largely absent from the documents are social scien-
tists. For example, none of the documents identied a need for partici-
pation by or collaboration with social scientists. de Graeff et al. (2019)
noted the absence of social science and humanities scholars in shaping
the academic debate on gene editing more broadly. Yet the involvement
of social scientists will be critical for addressing global problems such as
those in conservation and invasive species management (Shah, 2020).
They will play an important role in helping address complex governance
issues, such as engagement (including that of Indigenous peoples), re-
sponsibility and ethics.
Interestingly, the principles reveal contrasting motivations for
engagement that may be in conict. The rst and second principles are
concerned with inclusive, broad and empowered engagement as well as
securing public acceptance. In the broader eld of gene drive, Leding-
ham and Hartley (2021) note there is often considerable slippage from
the ambitions of engagement as empowering and transformative and the
more reductive framing of engagement as a tool to gain a social license
and secure acceptability. Indeed, the concept of ‘social licence to oper-
ate’ has been found to be particularly problematic for gene drive (Del-
borne et al., 2020). In order for governance principles to provide a
framework for decisions, clarity on the matter of engagement is needed
to avoid such slippage and make clear how tensions will be managed.
There may also be a danger of conict of interest and ‘governance
capture’ by developers and supporters. For example, Pohl et al. (2010)
highlight why approaches to tackling drought and soil degradation have
failed to work in practice. This is because they failed to take into account
relationships of mistrust between farmers and government authorities
and other knowledge regimes that constitute different ways of living/-
doing/being. They highlight concerns about co-production processes
becoming ‘hijacked’ by powerful interests including local elites, com-
panies and government agencies. As engagement and knowledge
co-production needs are further dened in conservation gene drive, it
will be important to ensure a broad range of actors are involved in
dening the processes through which engagement and co-production
will take place, not just participating in engagement that has been
dened by those in power.
In addition to the matter of engagement, our ndings reveal other
areas of interest and gaps that are surprising given the application
domain of conservation and invasive species management. For example,
given the explicit focus on conservation applications, it was surprising to
see very limited mention of the precautionary principle, which is a key
principle in biotechnology governance, particularly for GMOs. Further,
the conservation community is diverse in terms of its worldviews. Gene
drive applications in global health raise similar hopes and concerns
among stakeholders. For example, stakeholder hopes are that gene drive
mosquitoes will eradicate malaria while concerns are focused on envi-
ronmental protection, the economy and governance (Hartley et al.,
2021b). However, the conservation community is unlikely to speak with
a united voice. For example, Sandbrook et al. (2019) highlight the
diverse range of positions in conservation and it is not yet clear how
these actors will respond to potential gene drive applications in their
eld. This diversity is not recognized in the documents yet any
engagement activity will need to be sensitive to the diverse range of
actors in conservation who may hold different and conicting values.
Governance frameworks for emerging technologies must be attentive
to the technology’s social, ethical and political realities (Marchant and
Tournas, 2019). Principles are a means to do this. However, these
principles are emergent – they present a collection of visions of an ideal
state of governance from different perspectives – and they may not sit
well together. Responsibility is dened quite differently across the
documents, with some framing responsibility as governance that antic-
ipates transboundary effects, others emphasizing deliberation, and
others emphasizing continued technological innovation. The ethical
imperative for gene drive applications in conservation is not as clear as it
is for applications in global health and the increased range of actors
combined with the contested nature of conservation may result in a
more challenging governance debate. However, exposing the principles,
as we have done here, allows for stakeholders to examine and debate the
merits of the different perspectives as governance frameworks are pin-
ned down by a variety of actors, including frameworks for the funding,
regulation and potential deployment of gene drive organisms. The
principles are persuasive rhetorically, but are they identiable in prac-
tice? The next step is to think about who the principles are intended for,
how they will be used and embedded into practice and policy, and how
they can be made meaningful. These principles are emergent and
considerable denition is needed before a tangible and clear governance
framework could embody them.
Declaration of Competing Interest
The authors declare that they have no known competing nancial
interests or personal relationships that could have appeared to inuence
the work reported in this paper.
Acknowledgements
This research was funded through a QUEX Accelerator Grant
administered by the University of Exeter, UK and the University
Queensland, Australia. The research data supporting this publication are
provided within this paper.
References
Akbari, O.S., Bellen, H.J., Bier, E., Bullock, S.L., Burt, A., Church, G.M., Cook, K.R.,
Duchek, P., Edwards, O.R., Esvelt, K.M., Gantz, V.M., Golic, K.G., Gratz, S.J.,
Harrison, M.M., Hayes, K.R., James, A.A., Kaufman, T.C., Knoblich, J., Malik, H.S.,
Matthews, K.A., O’Connor-Giles, K.M., Parks, A.L., Perrimon, N., Port, F., Russell, S.,
Ueda, R., Wildonger, J., 2015. Safeguarding gene drive experiments in the
laboratory. Science 349, 927–929.
Allot, A., 2021, Genetically-modied possums and all-in-one trapping machines: funding
for new predator-free studies, Stuff, New Zealand.
Annas, G.J., Beisel, C.L., Clement, K., Crisanti, A., Francis, S., Galardini, M., Galizi, R.,
Grunewald, J., Immobile, G., Khalil, A.S., Muller, R., Pattanayak, V., Petri, K.,
Paul, L., Pinello, L., Simoni, A., Taxiarchi, C., Joung, J.K., 2021. A Code of Ethics for
Gene Drive Research. CRISPR J. 4, 19–24.
Ansell, C., Torng, J., 2016. Introduction: theories of governance. In: Ansell, C.,
Torng, J. (Eds.), Handbook on Theories of Governance. Edward Elgar Publishing,
Cheltenham.
Arnstein, S.R., 1969. A ladder of citizen participation. J. Am. Inst. Plan. 35, 216–224.
Arvin, M., 2019. Possessing Polynesians: The Science of Settler Colonial Whiteness in
Hawai’i and Oceania. Duke University Press,, Durham.
Barnhill-Dilling, S.K., Delborne, J.A., 2021. Whose intentions? What consequences?
Interrogating “Intended Consequences” for conservation with environmental
biotechnology. Conserv. Sci. Pract. 3 (4), e406.
Barnhill-Dilling, S.K., Delborne, J.A., 2019. The genetically engineered american
chestnut tree as opportunity for reciprocal restoration in haudenosaunee
communities. Biol. Conserv. 232, 1–7.
Barnhill-Dilling, S.K., Rivers, L., Delborne, J.A., 2020. Rooted in recognition: indigenous
environmental justice and the genetically engineered american chestnut tree. Soc.
Nat. Resour. 33, 83–100.
Barnhill-Dilling, S.K., Serr, M., Blondel, D.M., Godwin, J., 2019. Sustainability as a
Framework for Considering Gene Drive Mice for Invasive Rodent Eradication.
Sustainability 11.
Benedict, M., D’Abbs, P., Dobson, S., Gottlieb, M., Harrington, L., Higgs, S., James, A.,
James, S., Knols, B., Lavery, J., O’Neill, S., Scott, T., Takken, W., Toure, Y.,
Containe, C.W.G.G., 2008. Guidance for contained eld trials of vector mosquitoes
engineered to contain a gene drive system: Recommendations of a scientic working
group. Vector-Borne Zoonot 8, 127–166.
S. Hartley et al.
Environmental Science and Policy 135 (2022) 36–45
44
Benedict, M.Q., Burt, A., Capurro, M.L., De Barro, P., Handler, A.M., Hayes, K.R.,
Marshall, J.M., Tabachnick, W.J., Adelman, Z.N., 2018. Recommendations for
Laboratory Containment and Management of Gene Drive Systems in Arthropods.
Vector-Borne Zoonot 18.
Braun, V., Clarke, V., 2019. Reecting on reexive thematic analysis. Qual. Res. Sport,
Exerc. Health 11, 589–597.
Burall, S., 2018. Rethink public engagement for gene editing. Nature 555, 438–439.
Callaway, E., 2018. Controversial CRISPR ‘gene drives’ tested in mammals for the rst
time. Nature 559.
Claw, K.G., Anderson, M.Z., Begay, R.L., Tsosie, K.S., Fox, K., Garrison, N.A., 2018.
A framework for enhancing ethical genomic research with Indigenous communities.
Nat. Commun. 9 (1), 1–7.
Cohen-Shacham, E., Andrade, A., Dalton, J., Dudley, N., Jones, M., Kumar, C.,
Maginnis, S., Maynard, S., Nelson, C.R., Renaud, F.G., Welling, R., Walters, G., 2019.
Core principles for successfully implementing and upscaling Nature-based Solutions.
Environ. Sci. Policy 98, 20–29.
Convention on Biological Diversity, 2017, Report of the Ad Hoc Technical Expert Group
on Synthetic Biology. Conference of the Parties to the Convention on Biological
Diversity, 1–17.
Cornwall, W., 2019. Researchers embrace a radical idea: engineering coral to cope with
climate change. Science.
de Graeff, N., Jongsma, K.R., Johnston, J., Hartley, S., Bredenoord, A.L., 2019. The ethics
of genome editing in non-human animals: a systematic review of reasons reported in
the academic literature. Philos. Trans. R. Soc. B 374 (1772), 20180106.
Delborne, J.A., Kokotovich, A.E., Lunshof, J.E., 2020. Social license and synthetic
biology: The trouble with mining terms. J. Responsible Innov. 7 (3), 280–297.
Diagne, C., Leroy, B., Vaissi`
ere, A.C., Gozlan, R.E., Roiz, D., Jari´
c, I., Salles, J.-M.,
Bradshaw, C.J.A., Courchamp, F., 2021. High and rising economic costs of biological
invasions worldwide. Nature 592, 571–576.
Dumas, P., Fiat, S., Durbano, A., Peignon, C., Moutham, G., Ham, J., Gereva, S., Kaku, R.,
Chateau, O., Wantiez, L., De Ramon N’Yeurt, A., Adjeroud, M., 2020. Citizen
Science, a promising tool for detecting and monitoring outbreaks of the crown-of-
thorns starsh Acanthaster spp. Sci. Rep. 10.
Edwards, O., Brown, P., Tizard, M., Strive, T., Sheppard, A., 2017. Taking a responsible
approach to new genetic technologies for conservation. CSIRO,.
Emerson, C., James, S., Littler, K., Randazzo, F., 2017. Principles for gene drive research.
Science 358, 1135–1136.
Esvelt, K.M., Gemmell, N.J., 2017. Conservation demands safe gene drive. Plos Biol. 15,
e2003850.
Faber, N.R., McFarlane, G.R., Gaynor, R.C., Pocrnic, I., Whitelaw, C.B.A., Gorjanc, G.,
2021. Novel combination of CRISPR-based gene drives eliminates resistance and
localises spread. Sci. Rep. 11.
Fox, K., Rallapalli, K.L., Komor, A.C., 2020. Rewriting human history and empowering
indigenous communities with genome editing tools. Genes 11 (1), 88.
Gardiner, S.M., Fragni`
ere, A., 2018. The tollgate principles for the governance of
geoengineering: Moving beyond the Oxford principles to an ethically more robust
approach. Ethics, Policy Environ. 21 (2), 143–174.
George, D.R., Kuiken, T., Delborne, J.A., 2019, Articulating ’free, prior and informed
consent’ (FPIC) for engineered gene drives. Proceedings of the Royal Society B:
Biological Sciences, 286(1917), 20191484.
Goldman, J.G., 2016, Harnessing the power of gene drives to save wildlife. Scientic
American, September issue, 14–19.
Golnar, A.J., Ruell, E., Lloyd, A.L., Pepin, K.M., 2020. Embracing dynamic models for
gene drive management. Sci. Soc. 39, 211–214.
Hagendijk, R., Irwin, A., 2006. Public deliberation and governance: engaging with
science and technology in contemporary Europe. Minerva 44, 167–184.
Hartley, S., Ledingham, K., Owen, R., Leonelli, S., Diarra, S., Diop, S., 2021a.
Experimenting with co-development: a qualitative study of gene drive research for
malaria control in Mali. Soc. Sci. Med. 276, 113850.
Hartley, S., Smith, R.D.J., Kokotovich, A., Opesen, C., Habtewold, T., Ledingham, K.,
Raymond, B., Rwabukwali, C.B., 2021b. Ugandan stakeholder hopes and concerns
about gene drive mosquitoes for malaria control: new directions for gene drive risk
governance. Malar. J. 20.
Hartley, S., Thizy, D., Ledingham, K., Coulibaly, M., Diabate, A., Dicko, B., Diop, S.,
Kayondo, J., Namukwaya, A., Nourou, B., Toe, L.P., 2019. Knowledge engagement in
gene drive research for malaria control. Plos Negl. Trop. D. 13, 1–5.
Høj, L., Levy, N., Baillie, B.K., Clode, P.L., Strohmaier, R.C., Siboni, N., Webster, N.S.,
Uthicke, S., Bourne, D.G., 2018. Crown-of-Thorns Sea Star Acanthaster cf. solaris Has
Tissue-Characteristic Microbiomes with Potential Roles in Health and Reproduction.
Appl. Environ. Microbiol. 84.
Holman, L., 2019, Evolutionary simulations of Z-linked suppression gene drives.
Proceedings of the Royal Society B 286.
Hughes, T.P., Barnes, M.L., Bellwood, D.R., Cinner, J.E., Cumming, G.S., Jackson, J.B.C.,
Kleypas, J., Leemput, I.A., van de, Lough, J.M., Morrison, T.H., Palumbi, S.R., Nes, E.
H., Van, Scheffer, M., 2017. Coral reefs in the Anthropocene. Nature 546, 82–90.
https://doi.org/10.1038/nature22901.
Hudson, M., Mead, A.T.P., Chagn´
e, D., Roskruge, N., Morrison, S., Wilcox, P.L., Allan, A.
C., 2019. Indigenous perspectives and gene editing in Aotearoa New Zealand. Front.
Bioeng. Biotechnol. 7, 70.
IPBES, 2019. Global Assessment Report on Biodiversity and Ecosystem Services of the
Intergovernmental Science (e.a.E). In: Brondizio, E.S. (Ed.), Policy Platform on
Biodiversity and Ecosystem Services. IPBES Secretariat, Bonn, Germany.
Jasanoff, S., 2004. States of Knowledge: The Co-production of Science and the Social
Order. Routledge,.
Kachel, N., 2018, Gene drive technology: A new hope in the ght against feral cats.
CSIROscope.
Kaebnick, G.E., 2021. Does gene editing in the wild require broad public deliberation?
Hastings Cent. Rep. 51, S34–S41.
Kamya, P.Z., Byrne, M., Mos, B., Hall, L., Dworjanyn, S.A., 2017, Indirect effects of ocean
acidication drive feeding and growth of juvenile crown-of-thorns starsh,
Acanthaster planci. Proceedings of the Royal Society B 284.
Kelsey, A., Stillinger, D., Binh Pham, T., Murphy, J., Firth, S., Carballar-Lejarazú, R.,
2020. Global governing bodies: a pathway for gene drive governance for vector
mosquito control. Am. J. Trop. Med. Hyg. 103, 976–985.
Ledingham, K., Hartley, S., 2021. Transformation and slippage in co-production
ambitions for global technology development: The case of gene drive. Environ. Sci.
Policy 116, 78–85.
Leitschuh, C.M., Kanavy, D., Backus, G.A., Valdez, R.X., Serr, M., Pitts, E.A.,
Threadgill, D., Godwin, J., 2018. Developing gene drive technologies to eradicate
invasive rodents from islands. J. Responsible Innov. 5, S121–S138.
Lester, P.J., Bulgarella, M., Baty, J., Dearden, P.K., Guhlin, J., Kean, J.M., 2020. The
potential for a CRISPR gene drive to eradicate or suppress globally invasive social
wasps. Sci. Rep. 10.
Littler, K., 2021. From principles to principled action: What ethical principles ought to
govern gene drive research? Gene Drive Res. Forum, “Unsettled Ethic-.-. Issues Gene
Drive Res. ”. Session 4, found at. 〈https://www.youtube.com/watch?v=5tPnIOE
Xw4M&ab_channel=GeneConveneGlobalCollaborative〉.
Long, K.C., Alphey, L., Annas, G.J., Bloss, C.S., Campbell, K.J., Champer, J., Chen, C.-H.,
Choudhary, A., Church, G.M., Collins, J.P., Cooper, K.L., Delborne, J.A., Edwards, O.
R., Emerson, C.I., Esvelt, K., Weiss Evans, S., Friedman, R.M., Gantz, V.M., Gould, F.,
Hartley, S., Heitman, E., Hemingway, J., Kanuka, H., Kuzma, J., Lavery, J.V., Lee, Y.,
Lorenzon, M., Lunshof, J.E., Marshall, J.M., Messer, P.W., Montell, C., Oye, K.A.,
Palmer, M.J., Papathanos, P.A., Paradkar, P.N., Piaggio, A.J., Rasgon, J.L., Raˇ
si´
c, G.,
Rudenko, L., Saah, J.R., Scott, M.J., Sutton, J.T., Vorsino, A.E., Akbari, O.S., 2020.
Core commitments for eld trials of gene drive organisms. Science 370.
Marchant, G., Tournas, L., 2019. Filling the governance gap: International principles for
responsible development of neurotechnologies. AJOB Neurosci. 10, 176–178.
Min, J., Smidler, A.L., Najjar, D., Esvelt, K.M., 2018. Harnessing gene drive.
J. Responsible Innov. 5, S40–S65.
NASEM, 2016. Gene drives on the horizon: advancing science, navigating uncertainty,
and aligning research with public values. The National Academies Press,,
Washington, D.C.
NIH, 2021. Exceptional Technology and Research Advisory Committee: Gene Drives in
Biomedical Research Report. National Institutes of Health,.
OECD, 2019. Recomm. Counc. Artif. Intell. 0449.
Oye, K.A., Esvelt, K., Appleton, E., Catteruccia, F., Church, G., Kuiken, T., Lightfoot, S.B.
Y., McNamara, J., Smidler, A.L., Collins, J.P., 2014. Regulating gene drives. Science
345, 626–628.
Palmer, S., Mercier, O.R., King-Hunt, A., 2020. Towards rangatiratanga in pest
management? M¯
aori perspectives and frameworks on novel biotechnologies in
conservation. Pacic Conservation Biology. CSIRO,.
Piaggio, A.J., Segelbacher, G., Seddon, P.J., Alphey, L., Bennett, E.L., Carlson, R.H.,
Friedman, R.M., Kanavy, D., Phelan, R., Redford, K.H., Rosales, M., Slobodian, L.,
Wheeler, K., 2017. Is it time for synthetic biodiversity conservation? Trends Ecol.
Evol. 32, 97–107.
Pohl, C., Rist, S., Zimmermann, A., Fry, P., Gurung, G.S., Schneider, F., Speranza, C.I.,
Kiteme, B., Boillat, S., Serrano, E., Hadorn, G.H., Wiesmann, U., 2010. Researchers’
roles in knowledge co-production: experience from sustainability research in Kenya,
Switzerland, Bolivia and Nepal. Sci. Public Policy 37, 267–281.
Rabitz, F., 2019. Gene drives and the international biodiversity regime. RECIEL 28,
339–348.
Rands, M.R.W., Adams, W.M., Bennun, L., Butchart, S.H.M., Clements, A., Coomes, D.,
Entwistle, A., Hodge, I., Kapos, V., Scharlemann, J.P.W., Sutherland, W.J., Vira, B.,
2010. Biodiversity Conservation: Challenges Beyond 2010. Science 329, 1298–1303.
Rayner, S., Heyward, C., Kruger, T., Pidgeon, N., Redgwell, C., Savulescu, J., 2013. The
Oxford Principles. Clim. Change 121, 499–512.
Redford, K.H., Adams, W., Carlson, R., Mace, G.M., Ceccarelli, B., 2014. Synthetic
biology and the conservation of biodiversity. Oryx 48, 330–336.
Redford, K.H., Brooks, T.M., Macfarlane, N.B.W., Adams, J.S., 2019, Genetic frontiers for
conservation: An assessment of synthetic biology and biodiversity conservation.
Technical assessment. IUCN, Gland, Switzerland.
Reynolds, J.L., 2020. Governing New Biotechnologies for Biodiversity Conservation:
Gene Drives, International Law, and Emerging Politics. Glob. Environ. Polit. 20,
28–48.
Royal Society Te Ap¯
arangi, 2017, The use of gene editing in pest control, New Zealand.
Rudenko, L., Palmer, M.J., Oye, K., 2018. Considerations for the governance of gene
drive organisms. Pathog. Glob. Health 112, 162–181.
Rudgard, O., 2021, Genetic modication could be used to combat invasive craysh, The
Telegraph, UK.
Sandbrook, C., Fisher, J.A., Holmes, G., Luque-Lora, R., Keane, A., 2019. The global
conservation movement is diverse but not divided. Nat. Sustain. 2, 316–323.
Shah, H., 2020. Global problems need social science. Nature 577 (7789), 295–296.
Shaw, S., Elston, J., Abbott, S., 2006. Comparative analysis of health policy
implementation: The use of documentary analysis. Policy Stud. 25, 259–266.
Sovacool, B.K., Axsen, J., Sorrell, S., 2018. Promoting novelty, rigor, and style in energy
social science: Towards codes of practice for appropriate methods and research
design. Energy Res Socilal Sci 45, 12–42.
Stelmach, A., Nerlich, B., Hartley, S., 2022. Gene drives in the U.K., U.S., and Australian
press (2015-2019): How a new focus on responsibility is shaping science
communication. Sci. Commun.
S. Hartley et al.
Environmental Science and Policy 135 (2022) 36–45
45
Stirling, A., 2008. “Opening Up” and “Closing Down”: power, participation, and
pluralism in the social appraisal of technology. Sci., Technol., Hum. Values 33,
262–294.
Stirling, A., 2010. From enlightenment to enablement: opening up choices for
innovation. The Innovation for Development Report. Palgrave Macmillan,, London,
pp. 199–210.
Sustainability Council of New Zealand, 2018, A Constitutional Moment: Gene Drive and
International Governance, New Zealand.
Taitingfong, R., 2019. Islands as laboratories: Indigenous knowledge and gene drives in
the Pacic. Hum. Biol. 91, 179.
Taitingfong, R., Ullah, A., 2021, Empowering Indigenous Knowledge in Deliberations on
Gene Editing in the Wild, in Gene Editing in the Wild: Shaping Decisions through
Broad Public Deliberation, ed. Michael K. Gusmano et al., special report, Hastings
Center Report 51, no. 6: S74-S84.
TallBear, K., 2013. Native American DNA: Tribal Belonging and the False Promise of
Genetic Science, 1 edition.,. Univ Of Minnesota Press,, Minneapolis, MN.
Thizy, D., Coche, I., de Vries, J., 2020. Providing a policy framework for responsible gene
drive research: an analysis of the existing governance landscape and priority areas
for further research. Wellcome Open Res. 5.
Trouwborst, A., McCormack, P.A., Camacho, E.M., 2020. Domestic cats and their impacts
on biodiversity: A blind spot in the application of nature conservation law. People
Nat. 2, 235–250.
Tsosie, K.S., Claw, K.G., 2020. Indigenizing science and reasserting Indigeneity in
research. Hum. Biol. 91 (3), 137–140.
Ulnicane, I., Knight, W., Leach, T., Stahl, B.C., Wanjiku, W.-G., 2020. Framing
governance for a contested emerging technology:insights from AI policy. Policy Soc.
40, 158–177.
Weiss Evans, S., Palmer, M.J., 2018. Anomaly handling and the politics of gene drives.
J. Responsible Innov. 5, S223–S242.
Wynne, B., 1992. Uncertainty and environmental learning: Reconceiving science and
policy in the preventive paradigm. Glob. Environ. Change 2, 111–127.
S. Hartley et al.