Literature Review on Genetic Use Restriction

Technologies
April 2006

By Tony Szumigalski, Ph.D.

403 Gilmore Ave., Winnipeg, Manitoba
(204)663-1642
[email protected]

Abstract
The purpose of this paper is to conduct a literature review to survey current technical and
related literature on genetic use restriction technologies (GURTs). Topics covered
include the nature of current or proposed GURTs, the potential benefits of GURTs, the
risks and potential costs associated with GURTs, and a comparative study between
GURTs and hybrid seed technologies. Since the initial patent for terminator technology
was granted in 1998, the issue of GURTs has been complex and divisive, with different
stakeholders embracing widely opposing views. There are several strong arguments that
have been put forward, both in favour and against the implementation of these types of
technologies. Potential benefits include intellectual property rights protection, stimulation
of private research and development, genetic diversity enhancement, transgene
containment and production purposes. Potential risks and costs associated with GURTs
include outcrossing of terminator genes, reduced access and increased cost of genetic
material for breeders, greater necessity of regulating and field monitoring of new GURT
technologies, liability for environmental damage, health risks, increased cost of seeds for
farmers, greater corporate control over agriculture, and a further narrowing of agrobiodiversity. Until the results of peer-reviewed research on the environmental, social,
economic and political impacts of GURTs are publicly available, it is recommended that
a precautionary approach regarding implementation of GURTs be followed.

Table of Contents
Abstract ................................................................................................................................ i
Table of Contents................................................................................................................ ii
1. Introduction..................................................................................................................... 1
2. Current status of genetic use restriction technologies..................................................... 1
2.1. Trait based GURTs ............................................................................................................... 1
2.2. Variety based GURTs (sterile seed technology)................................................................... 2

3. Benefits of GURTs ......................................................................................................... 3

3.1 Private seed companies (plant breeders) .............................................................................. 4
3.2 Farmers.................................................................................................................................. 5
3.3 Governments ......................................................................................................................... 6
3.4 Society ................................................................................................................................... 6

4. Risks of GURTs.............................................................................................................. 7
4.1. Risks of transgene escape.................................................................................................... 7
4.2. Other risks and costs of GURTs ........................................................................................... 8
4.2.1 Private seed companies (plant breeders) ....................................................................... 8
4.2.2 Farmers ........................................................................................................................... 8
4.2.3 Governments................................................................................................................... 9
4.2.4 Society............................................................................................................................. 9

5. Comparison of GURTs and hybrid seed technology .................................................... 10

6. Summary and conclusions ............................................................................................ 11
7. References..................................................................................................................... 12
Appendices........................................................................................................................ 14

ii

iii

1. Introduction
The purpose of genetic use restriction technologies (GURTs) is to restrict access to
genetic materials and their associated traits. This type of technology was developed to
ensure that new seed varieties could be protected against unauthorized use. Although
originally patented as a technology protection system for arable crops (United States
Patent and Trademark Office 1998), this type of system could potentially be applied to
any organism type, such as trees, livestock or fish (Visser et al. 2001). The topic of
GURTs has been a very controversial one, with strongly divergent stakeholders, since the
first patent for this technology was granted in 1998 (e.g., Service 1998; Mahajan 1999;
Pendleton 2004). Strong arguments can be made both in favour and against the utilization
of these types of technologies, depending on socio-economic, political and environmental
context.
The purpose of this paper is to conduct a literature review to survey current technical and
related literature on the complex issue of GURTs. This information will be used by the
Canadian Foodgrains Bank to prepare a discussion guide for further debate on the merits
and risks of GURTs for farming, both in developing countries and in Canada. The main
body of this literature review is divided into several different sections. In Section 2, the
nature of current or proposed genetic use restriction technologies, with particular
attention to sterile seed technologies, is reviewed. In Section 3, the potential benefits of,
and those who might benefit from GURTs, are investigated. In contrast, the risks and
potential costs associated with GURTs are examined in Section 4. In Section 5, a
comparative study is made between GURTs and hybrid seed technologies, with emphasis
from the point of view of their application in developing countries by small farmers.
Finally, Section 6 represents a brief summary of key points and major findings from this
review, and Section 7 is the bibliography.

2. Current status of genetic use restriction technologies

There are currently two main classes of GURTs: trait based and variety based GURTs
(Visser et al. 2001; Eaton et al. 2002; Pendleton 2004). Trait based technologies (TGURTs) regulate the expression of a particular trait; whereas variety based technologies
(V-GURTs) restrict the use of an entire variety by blocking its reproduction. There is
often a blur in distinction between the two types of GURTS and the molecular
mechanism employed in both is very similar (Federation of German Scientists 2006).
Although this review will focus on V-GURTs, both types of technologies will be
discussed below.

2.1. Trait based GURTs

In the case of T-GURTs, one or more genes conferring a single trait are switched on or
off through application of chemical inducers (Visser et al. 2001; Pendleton 2004).
Therefore, T-GURTs are not intended to affect the viability of seeds, which is in contrast
to V-GURTs, which result in sterile seeds. Traits of interest that could be controlled by
T-GURTs include male sterility, pest resistance, stress tolerance, nutrient production,
seed germination or flower development (Gupta 1998; Pendleton 2004). The goal of this
technology is to protect intellectual property (i.e., the value-added transgenic trait of

Page 1 of 21

interest) of plant breeders in newly developed varieties by restricting access through a

biological mechanism (Eaton et al. 2002). These traits could be activated when needed by
induction chemicals. For example, insecticidal genes (e.g., Bt) under the control of an
inducible promoter could remain inactivated until an insect pest outbreak justified the
application of a chemical to induce the formation of gene products toxic to insects. The
inducer chemical would, of course, be under the control and licensing of the seed
company; however, the ultimate trigger of this technology could be under the control of
producers (Pendleton 2004).

2.2. Variety based GURTs (sterile seed technology)

The original patent for V-GURTs (US Patent 5,723,765) entitled Control of plant gene
expression was granted jointly to the United States Department of Agriculture (USDA)
and Delta and Pine Land (D&PL) Company of Mississippi in March 1998 (United States
Patent and Trademark Office 1998). See Appendix 1 for an abstract of this patent.
Although this patented technology was originally developed for tobacco and cotton, it
could potentially be applied to all seed-propagated crops (Lehmann 1998). This
technology results in a change in the genetic makeup of a plant cell, whereby plants
regenerated from this cell will develop non-viable seeds, which will not germinate in the
next generation. Therefore, farmers who use seeds protected by this technology will be
able to grow and harvest a first crop, but will not be able to save (brown bag) seeds
from this crop to plant in the future. Although, there were originally plans for a corporate
merger between D&PL and Monsanto following this patent, this merger eventually failed
(Pilger 2002), largely in response to public pressure against Monsanto not to introduce
this germination control technology (Masood 1998; Niiler 1999). However, the USDA
decided to pursue the commercialization of this technology (Kaiser 2000).
Briefly, this system, also known as terminator technology, is based on the transfer of a
combination of three genes (Gupta 1998; Lehmann 1998; Pendleton 2004). These genes
are: 1) a gene coding for a toxic substance (terminator or lethal gene), which is linked to a
blocking sequence preventing the activation of the terminator gene, 2) a recombinase
gene (CRE/LOX) containing the information for a protein which cuts the blocking
sequence linked to the toxic gene, and 3) a repressor gene with the code for a protein
which suppresses the recombinase gene and which is controlled by an external stimulus
(Figure 1; see Appendix 2 for more detailed diagram). Without the application of an
external stimulus (i.e., chemical trigger such as the antibiotic tetracycline) the repressor
gene is normally switched on; therefore, the recombinase gene is switched off so that the
blocking sequence of the terminator gene remains intact. Thus, under normal conditions
(absence of chemical trigger) the crop remains fertile. However, once the chemical
trigger is released, the repressor gene is switched off allowing for the recombinase gene
to be switched on, which in turn removes the blocking sequence from the terminator
gene, resulting in lethal gene expression. Toxins produced by the activated terminator
gene destroy the embryo, thereby rendering the seeds sterile. In this way, seeds could be
produced and increased commercially in the absence of the chemical trigger, but when
sold to farmers the seeds would be treated with the chemical. All other aspects of plant
growth would remain unaffected, because the toxic effects stimulated by chemical
treatment only occur during the later stages of embryo development, thus not adversely
affecting final yields (Lehmann 1998).
Page 2 of 21

Figure 1: Simplified V-GURTs activity

Besides the mechanism described above for the original patent, there are a number of
other variations of the same basic theme that can be applied to interfere with reproduction
in V-GURTs. The system described above pertains mainly to pure line seed production
in self-pollinated crops; however, in the case of hybrid seed production an alternate
strategy utilizing different gene components in each hybrid parent could be used (Gupta
1998; Lehmann 1998; Pendleton 2004). In this system, the first hybrid generation (grown
by the farmer) would be sterile because the lethal gene and recombinase would be
brought together in this generation (Gupta 1998) and a chemical trigger would not be
necessary (Lehmann 1998). In another system, tetracycline or another chemical trigger
could also be used to inactivate the terminator trait instead of being used to activate the
trait (Visser et al. 2001; Pendleton 2004). Another strategy for V-GURTs pertains to
vegetatively reproducing crops (e.g., roots, tubers and some ornamentals), wherein,
unwanted growth during storage can be prevented by inactivating a certain gene, but
growth can restored when needed by activating a second gene (Visser et al. 2001).
Indeed, several companies have been pursuing a number of different patents on
technology that is similar to the original terminator technology (Kaiser 2000).

3. Benefits of GURTs
There are several potential benefits, costs and risks associated with GURTs (Table 1).
This section will focus on the benefits, whereas the following section (Section 4) will
focus on the risks associated with the deployment of GURTs. Potential benefits include
intellectual property rights protection, stimulation of private research and development,
genetic diversity enhancement, transgene containment and production purposes (Gupta
1998; Lehman 1998; Visser et al. 2001; Eaton et al. 2002; Goeschl and Swanson 2003;
Lence et al. 2005). The degree of potential benefits derived from GURTs depends on
social group (i.e., private companies, farmers, government or society in general). Positive
impacts of GURTs on each group will be discussed below. It should be noted that these
groups are not necessarily mutually exclusive.

Page 3 of 21

Table 1: Genetic use restriction technology (GURT): potential economic benefits, costs
and risks (Modified from Eaton et al. 2002).
.

Benefits

Costs

Risks
Misuse of monopoly
powers by breeders

Farmers

Increased productivity
from improved genetic
inputs due to increased
research and
development (R&D)
investment

Increased input
costs from seed
purchase

Breeders
(especially private
sector)

Increased share of
research benefits from
new products

Increased cost for

access to genetic
resources of other
breeders

Reduced seed
security and access
to genetic
improvements
(marginalized
farmers)

Reduced investment
requirements in breeding
Governments

Complementary
R&D investment
requirements
Establishment and
Fewer enforcement costs enforcement of new
for plant variety protection regulatory
(PVP)
requirements

Society

Increased agricultural
productivity

Reduced genetic
diversity in fields

3.1 Private seed companies (plant breeders)

It has been suggested that the absence of intellectual property rights results in diminished
private research and development (R&D) in plant breeding (Goeschl and Swanson 2003;
Pendleton 2004; Lence et al. 2005). For example, more private research has traditionally
been conducted on crops for which hybrids are feasible (e.g., corn and sorghum)
compared to those for which they are not (e.g., wheat and rice) (Goeschl and Swanson
2003). Therefore, GURTs represent a novel mechanism for capturing returns from
innovation in the plant breeding industry, in a similar manner to more conventional
hybridizing techniques. The GURT mechanism greatly improves the plant breeders
capacity for rent capture, potentially increasing private investment into agricultural R&D
and, hence, a higher rate of innovation in the plant breeding industry (Goeschl and
Swanson 2003). Breeding companies hope to protect their investments in improved
varieties, thus, GURTs may present a better form of insurance (i.e., a biological one)
against the free use of genetic innovations than patents, plant breeders rights or licenses
(Visser et al. 2001; Pendleton 2004; Burk 2004). GURTs would allow better enforcement
of property rights (Lence et al. 2005). Apart from the sterile seed technology of VPage 4 of 21

GURTs, it is also possible that T-GURTs protecting value added traits in newly released
commercial varieties could be applied to virtually all crops (Visser et al. 2001). Plant
breeders and seed companies, thus stand to make substantial intellectual and financial
gains through implementation of GURTs.
The potential for transgene escape may also be reduced through sterile seed technology
(Gupta 1998; Visser et al. 2001; Eaton et al. 2002). This would be beneficial for seed
companies because it would decrease the probability of corporate liability for
environmental contamination or health risks due to escaped transgenes (Pendleton 2004),
but would also reduce the chances of competitors or farmers accessing proprietary
genetic material through volunteer or feral crop plants.
To police the unauthorized use of agro-biotechnology, seed companies must currently
send agents out into farmers fields, which along with ensuing lawsuits, can be a costly
process (Burk 2004). With enhanced transgene containment through GURTs, seed
corporations would save on costs of monitoring farmers fields for any unauthorized used
of copyrighted genetic material or transgene escapes that must be mitigated.

3.2 Farmers
Potential gains for farmers through implementation of GURTs may be related to
improved yields as a result of increased R&D on crop varieties by private seed companies
(Lehmann 1998; Eaton et al. 2002; Goeschl and Swanson 2003). From arguments already
stated in the sub-section above, it can be surmised that plant breeders will have greater
motivation to develop new and improved varieties of crops for which hybrids are
impracticable. For instance, with application of GURTs, it is expected that R&D will
increase for self-pollinated crops such as wheat, rice and cotton (Visser et al. 2001).
Farmers, thus, may profit in the long-term from these innovations because more
productive varieties will become available as breeding efforts increase. For example, the
vast majority of improved varieties have been from hybridized crops, with an average
annual yield growth of 2.18% over the long-term for hybrid crops compared to a value of
1.58% for non-hybrid crops in developed countries (Goeschl and Swanson 2003). Thus,
for many farmers, the surplus of increased yields compensates for having to buy new
hybrid seeds every year (Lehmann 1998). The utilization of GURTs potentially could
bring non-hybrid yield increases in line with those of hybrids. Furthermore, incentives to
breed new varieties may enhance genetic diversity in many important crops, thereby
providing further long-term benefits associated with biodiversity (e.g., pest resistance) to
farmers (Lehmann 1998).
Apart from long-term yield and biodiversity effects, use of GURTs may offer some shortterm practicable applications for farmers as well. Terminator technology could
effectively eliminate the problem of genetically modified (GM) crop volunteers in
farmers fields (Pilger 2002) and reduce potential for outcrossing with, and increasing the
fitness of, weedy relatives (Gupta 1998; Visser et al. 2001). Controlling GM volunteer
weeds such as herbicide resistant crops can add expense to farmers operating costs, since
alternative, often more expensive, herbicides may be required for their control. Another
possible agronomic benefit associated with terminator technology is that, because the
seeds are sterile in V-GURTs, the problem of crop sprouting would not be an issue with
this type of technology (Pilger 2002). In addition, farmers may want to restrict the
Page 5 of 21

expression of a trait only to a specific phase in the development of the plants or animals
or during biotic or abiotic stress (Visser et al. 2001). For instance, T-GURTs may allow a
stress response to occur only when really needed, because stress responses can require a
great deal of resources and may diminish the quality of the processed product. An
example of a stress response is a response (e.g., chemical production) to insect pest attack
in plants. Alternatively, V-GURTs may be utilized to manage successful reproduction of
farm animals in order to maintain the integrity of adapted crossbreeds, produced from
mating between local and high-yielding commercial breeds (Visser et al. 2001).

3.3 Governments
Governments may benefit from GURTs through reduced investment requirements in
breeding and fewer enforcement costs for plant variety protection (Eaton et al. 2002;
Pendleton 2004). Governments could, thus, use GURTs as justification to decrease
funding to agriculture R&D and biosafety/copyright infringement enforcement programs.
Governments are often under political pressure to reduce spending and cut taxes,
therefore, political points could be gained by reducing expenses in these areas. If
implementation of GURTs results in yield gains and benefits to farmers, then
governments can gain politically with policies that support GURTs.

3.4 Society
Society in general, which includes consumers and the environment, may derive some
benefits from GURTs. Society, as a whole, may benefit from GURTs as a result of
increased agricultural productivity (Eaton et al. 2002). Pendleton (2004) stressed the
importance of the private sectors role in providing adequate food for the worlds
burgeoning population. Arguments can be made that GURTs will foster an environment
of increased innovation, which is needed to develop technology capable of increasing
yields to meet global demands (Goeschl and Swanson 2003; Lence et al. 2005). However,
forecast model simulations conducted by Goeschl and Swanson (2003) suggested that the
most advanced countries stand to benefit the most in terms of productivity gains from
GURTs, while less advanced countries stand to gain the least over the medium to long
term (i.e., over 20 years). Historically, the general public (especially in developed
countries) has benefited greatly from inflated agricultural productivity. For instance, the
average Canadian now spends less than 10% of his or her income on food (Kirkpatrick
and Tarasuk 2003), and this cheap food policy can, arguably, be attributed to overproduction of commodities, which drives down prices, in addition to the effects of
subsidies. This was confirmed in a model investigating the impacts of intellectual
property protection (IPP) in the seed industry by Lence et al. (2005), who found that
while generally producers might lose from innovations, increased benefits to consumers
outweighed producer losses resulting in total overall welfare increases with IPP.
There are also believed to be environmental benefits associated with GURTs. Perhaps the
most often cited environmental benefit related to the application of GURTs is the
containment of potentially harmful transgenes (Gupta 1998; Visser et al. 2001; Pilger
2002; Pendleton 2004). Supposedly, GURTs could be used for the environmental
containment of entire transgenic varieties through use of V-GURTs or for specific
transgenes contained in transgenic varieties through use of T-GURTs (Visser et al. 2001).
T-GURTs could also be used to contain traits that might pose a health risk to consumers
Page 6 of 21

or farmers. In either case, biosafety risks should be reduced with GURTs; however, with
sterile seed technology, there is still a risk of GM pollen escaping and outcrossing with
other varieties or weedy relatives (Pendleton 2004). No research has yet to be published
on this topic.

4. Risks of GURTs
4.1. Risks of transgene escape
It should be noted that the risks of transgene escape associated with GURTs are poorly
understood, because there is, currently, no peer-reviewed published research on this
specific topic (Federation of German Scientists 2006). One of the most often cited
environmental risks associated with GURTs is the threat of outcrossing or gene flow of
terminator genes into crops or populations of wild relatives (Bhatia 1998; Lehmann
1998; Service 1998; Visser et al. 2001; Giovannetti 2003). The movement of genes
among crops and their wild relatives is possible through two mechanisms: dispersal in
viable pollen or in viable seeds (Daniell 2002; Schiemann et al. 2005). Because sterile
seed technology is designed to eliminate production of viable seeds, it is expected that
risks of transgene escape through the seed route will be very limited or impossible with
GURTs (Lehmann 1998; Rakshit 1998; Pendleton 2004). However, there are still
concerns of transgene escape via seeds. For instance, the 100% efficacy of tetracycline
(or some other chemical inducer) treatment on terminator seeds is questionable and the
recombinase gene (see Appendix 2) could remain inactive in some seeds (Giovannetti
2003). Considering that millions of terminator seeds would need to be treated with a
chemical inducer to render them sterile, it is not unreasonable to expect that at least a few
seeds will escape the effects of the chemical trigger. Although on a percentage basis the
number of unaffected seeds may be extremely small, on an absolute basis, this number
could still be potentially large, if hypothetically, sizable volumes of seeds were treated in
the same facility. Such escaped seeds would carry the complete genetic complement of
the V-GURT, and could go on to germinate to produce both pollen and more seeds
carrying the terminator technology trait. Costly fumigation treatments with the chemical
inducer to control possible escaped seeds and progeny may be required to mitigate
contaminated fields. Furthermore, if a GURT used a variation of the terminator
technology exhibiting negative control of a trait (i.e., the trait is expressed unless blocked
by an inducer - Visser et al. 2001; Pendleton 2004), then it is possible that terminator trait
could be blocked by related compounds that occur naturally or are applied intentionally.
Thus, utmost care must be made in selecting highly specific inducers for these types of
GURT systems (Visser et al. 2001).
Risks of outcrossing through pollen flow from GURTs appear to be greater than they are
through seed dispersal; since plants from chemically induced seeds are not inhibited from
producing pollen in the terminator technology. Gene escape through transgenic pollen
flow and cross-pollination is unavoidable in many of the worlds most important crops,
such as wheat, rice, corn, barley, sorghum and sugar beets (Giovannetti 2003). In
addition, spontaneous hybridization with wild relatives appears to be a common feature
of most crops (Ellstrand et al. 1999). Negative effects of outcrossing from V-GURT
varieties on viable seed yields of neighbouring crops are of special concern (Visser et al.
2001; Giovannetti 2003; Pendleton 2004). Pollen carrying a terminator trait could result
Page 7 of 21

in the sterility of the next generation of seeds, but this would not be realized until after
these seeds were planted. The reduced seed viability due to terminator pollen
contamination may be problematic to farmers who save seed to be planted the next year
or for seed growers who must provide high quality seed to customers. The magnitude of
these effects will depend on the outcrossing rate of the crop and on the distance between
the donor and acceptor plants (Visser et al. 2001). Outcrossing rates vary greatly between
crops, from less than 1% in most self-pollinated crops to 15-20% in strongly crosspollinated crops such as pigeon pea (Bhatia 1998). However, in some cases outcrossing
rates can also be high in self-pollinating crops. For example, Lawrie et al. (2006) found
that outcrossing rates could be up to 10% for some varieties of wheat in a greenhouse
study, and for a field study, Matus-Cadiz et al. (2004) found that outcrossing in wheat can
occur over a distance of several meters, but decreased rapidly with distance from the
pollen source. However, for insect pollinated crops (e.g., canola) outcrossing can occur
over several hundred meters (Giovannetti 2003). Although GURTs should minimize the
outcrossing risk of crops with related species (Gupta 1998; Pendleton 2004), there is still
concern that the terminator gene may be able to move around and transform within
agroecosystems through insects, birds and possibly soil bacteria (Giovannetti 2003). The
probability of outcrossing for GM trees and fish is particularly high because of the long
distance pollen dispersal in the former and high probability of escapes in the latter (Visser
et al. 2001). Technologies utilizing male sterility to contain transgenes could be useful in
preventing outcrossing of terminator genes and have already been commercialized in
herbicide resistant rapeseed (Daniell 2002).

4.2. Other risks and costs of GURTs

Besides the risk of transgene escape, there are several other potential costs and risks
associated with the application of GURTs (Table 1). The level of impact of these
technologies depends on the issue and social group of concern.

4.2.1 Private seed companies (plant breeders)

Breeders will have an increased cost of acquiring genetic resources from private breeders
(Eaton et al. 2002). With increased intellectual property protection and proprietary of
materials it will be very difficult to share resources amongst competing companies and
institutions. Thus there will be a reduced atmosphere of sharing genetic resources with
the implementation of GURTs and other property rights protection. With less cooperation
amongst breeders, there is a possibility of achieving fewer scientific accomplishments
because resources will have to be spread around within different companies that may
basically be trying to achieve similar goals. There is also a certain amount of risk that
companies must take in performing R&D on products, such as GURTs, that may not have
political or public approval in the end. There is also the risk of being held liable for
environmental contamination or adverse health effects related to terminator transgene
escape.

4.2.2 Farmers
GURTs may be potentially detrimental to farmers, especially poor farmers. First, there is
the possibility that traditional crops grown adjacent to those with sterile seed technology
may suffer reductions in viable seed due to pollen transfer from the latter to the former
Page 8 of 21

(Gupta 1998; Giovannetti 2003). Second, there would be increased input costs associated
with GURTs because farmers would have to purchase GURT seeds from the supplier
every year. This would be detrimental to farmers practice of saving seeds, which is
especially important, for poor farmers of developing countries (Gupta 1998; Lehmann
1998; Service 1998; Eaton et al. 2002). For example, about 90% of farmers in India save
seeds to be replanted (Bhatia 1998) and worldwide, greater than 1 billion people depend
on this practice (Service 1998). Sterile seed technology would also be detrimental to crop
genetic diversity in traditional farms, where farmers often breed and adapt local landraces
of crops and exchange seeds. Inputs of novel genetic material are of great benefit to
increase the genetic diversity of these locally adapted varieties; however, this practice
would not be possible with utilization of GURTs because novel genetic traits would be
under GURT control (Visser et al. 2001). Corporate concentration and increased controls
on farmers autonomy could threaten farmers food security. Richer countries and richer
farmers are likely to gain most of the benefits from this technology (Eaton et al. 2002;
Goeschl and Swanson 2003). Thus it appears that GURTs could further marginalize
already vulnerable and poor farmers.

4.2.3 Governments
With application of GURTs governments may have increased expenditures and
responsibilities (Eaton et al. 2002). Some governments might be obliged to provide
matching R&D investment to those of private companies. With development of GURTs,
governments may feel pressured to increase public funding for plant genetic resources
that are not copyright protected and have no genetic use restrictions. The uncertainty of
the containment of terminator transgenes may necessitate further regulations and
monitoring. Governments may also be liable for any ill environmental or health effects
due to GURTs. For example, the USDA (along with a private company D&PL) filed for
the first sterile seed patent (Appendix 1), so would therefore be partially responsible for
any damages accrued through the use of this technology. Allowing the commercialization
and release of terminator technology by governments might prove to be an unpopular
decision resulting in negative political ramifications (Service 1998; Niiler 1999).

4.2.4 Society
There are also potential risks posed to society by introduction of GURTs (Service 1998;
Visser et al. 2001; Eaton et al. 2002). Besides a risk of environmental contamination of
terminator transgenes, there is also a risk of reduced biodiversity in farmers fields due
further homogenization of crops. Although reduced crop diversity is already a serious
issue, it is believed that implementation of GURTs would further aggravate this problem,
because with sterile seed technology, farmers would not be have access to novel genetic
traits often utilized to increase agrobiodiversity at the local level (Visser et al. 2001). For
example, introgression of modern pearl millet varieties into locally adapted landraces in
India has been shown to increase genetic diversity of this crop (Yadav et al. 2000; vom
Brocke et al. 2002). A narrowing of the genetic pool for crop plants may result in
increased vulnerability to pest attack, which could destabilize yields and may result in
food shortages (e.g., the Irish potato famine). It is unclear how widespread adoption of
GURTs would impact local economies of countries (e.g., India), which rely heavily on
the exchange of seed for locally adapted varieties (Rakshit 1998). An over-reliance on

Page 9 of 21

biotechnology and associated inputs rather than on traditional knowledge, diversity and
ecological services to maintain production could be harmful to local agroecosystems.
While the advent Green Revolution has already initiated the erosion of local
agroecological capital (Visser 1998), it is expected that GURTs will exacerbate this
situation. With the need to purchase new seeds every year under GURTs, it is likely that
this technology would be detrimental to local cultures and economies of third world
countries where farmers typically save seed. . Similarly, risks may also arise from food
aid consisting of GURT seeds distributed for disaster relief, because relief grain supplies
are also often used as seed (Eaton et al. 2002). A further question is related to potential
impacts on human or animal nutrition associated with sterile seed technology. Since
terminator technology (in its original application) results in the destruction of the seed
embryo, there is a possibility that this technology could adversely affect the food quality
of harvested grain, because the embryo represents a nutritious portion of the seed with the
most protein. In the broader context of diet and health (e.g., type II diabetes) the issue of
food quality related to GURTs needs to be addressed.

5. Comparison of GURTs and hybrid seed technology

Hybrid technologies have several attributes which make them useful for plant breeding
and food production purposes (van Wijk 1994). These include a high yield potential,
relative ease of breeding in desirable traits, uniformity in maturity and size, and built-in
protection against multiplication. Therefore, hybrids have traditionally been considered
one of the most important incentives driving the development of a private seed industry
(van Wijk 1994; Goeschl and Swanson 2003; Pendleton 2004), and indeed, one can draw
parallels between hybrid technologies and GURTs in this respect. Hybridization has been
commercially applied to several major crops including maize, sorghum, rice and a
number of vegetables (Eaton et al. 2002).
There are several similarities as well as differences between hybrid seed technologies and
GURTs that can be noted. GURTs and hybrid seed technologies are similar in that they
both offer some form intellectual property protection to breeders and replanting of seeds
is not normally a viable option. Thus with both systems, farmers are discouraged from
saving seed and both may be used as use restriction technologies (Gupta 1998; Goeschl
and Swanson 2003; Pendleton 2004). However, for GURTs, these characteristics are
delivered in a more extreme form, whereby yield loss from replanting GURT seeds is
absolute and the reproduction of the seeds for breeding purposes is impossible (Goeschl
and Swanson 2003). In the case of hybrid seeds, yield reduction with replanting typically
results in a yield depression that is not absolute and, although F2 generations are inferior
to F1 ones, they can still be grown by the farmer and used for breeding purposes,
regardless of poor quality (Visser et al. 2001). It is possible that apomixis (seed from
vegetative reproduction and not from fertilization) could be applied to some hybrid crops,
thereby allowing farmers the possibility to produce seeds for the next growing season that
are the same as those of the parents (Visser 1998); however, apomixis is only known to
occur in a few crops and this type of technology is relatively undeveloped (Daniell 2002).
Currently, like GURTs, hybrid seeds make farmers dependent on purchasing external
seeds, which may result in a loss of autonomy and a financial burden on some farmers
(van Wijk 1994). In order to be economically feasible, hybrids generally must have 15 to

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20% higher yields than open-pollinated varieties to make up for the cost of purchase
(Lehmann 1998).
Hybrid F1 generations typically result in increased vigour or heterosis (Lehmann 1998;
Yadav et al. 2000); whereas this has yet to have been demonstrated for GURT seeds.
Instead, GURT crops are claimed to provide other agronomic traits (e.g., disease
resistance) that result in increased yields. It is possible, however, to utilize hybrid seed
technology in an alternate V-GURT strategy in which different gene components for the
terminator system are placed in each hybrid parent (Gupta 1998; Lehmann 1998;
Pendleton 2004). In this manner, the F1 generation would still produce sterile seed;
however, it is not known whether this type of system would result in increased vigour for
the first hybrid generation, because it is not known to what extent the genetic
backgrounds of the parents used for this cross differ in order to create effective heterosis
in the offspring. If the parents differ only by the inserted terminator genes, then heterosis
would be unlikely to be very great in the F1 generation of terminator seeds. If this were
the case, then the agronomic benefits associated with such a system would be derived
solely from transgenic traits and not from any hybrid vigour.
Although, all current GURTs are derived by genetic manipulation, this does not have to
be the case for hybrids. Therefore, while GM hybrids do exist, not all hybrids need be
GM crops, whereas all GURTs by definition must be GM. Thus non-GM hybrid crops
may not have the same market restrictions that GM crops currently have in some
jurisdictions. In short, it might be easier for farmers to sell hybrid crops compared to
crops grown using GURTs; however, this is unlikely to be an issue for small subsistence
farmers in third world countries where most grain is consumed locally and is not intended
for international markets.

6. Summary and conclusions

The issue of GURTs is complex, with different stakeholders embracing widely opposing
views. There are several strong arguments that have been put forward, both in favour and
against the implementation of these types of technologies. There are a number of
potential benefits, costs and risks associated with GURTs. Potential benefits include
intellectual property rights protection, stimulation of private research and development,
genetic diversity enhancement, transgene containment and production purposes. It seems
that private seed companies and consumers in general will gain the most, whilst farmers
(especially those with smaller holdings in third world countries) will gain the least as a
group from the adoption of GURTs. Environmental benefits associated with GURTs may
include enhanced containment of transgenes and fewer chemical inputs. However, one of
the most often cited environmental risks associated with GURTs is the threat of
outcrossing or gene flow of terminator genes into crops or populations of wild relatives.
Risks of outcrossing through pollen flow from GURTs appear to be greater than they are
through seed dispersal; however, terminator transgene escape through the latter route is
still probable. Other potential risks and costs associated with GURTs include reduced
access and increased cost of genetic material by breeders, greater necessity of regulating
and field monitoring of new GURT technologies, liability for environmental damage,
health risks, increased cost of seeds for farmers, greater corporate control over
agriculture, and a further narrowing of agro-biodiversity. Although there are many

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similarities between GURTs and hybrid seed technologies, it can be argued that impact of
former on small scale farmers would be more severe than that of the latter technology,
because restrictions associated with GURTs would be more absolute. Overall, it is very
difficult to assess the environmental, social, economic and political ramifications of
GURTs because there is a lack of peer-reviewed publications with novel research
addressing these issues. Until the results of this type of research are publicly available, it
is recommended that a precautionary approach regarding implementation of GURTs be
followed.

7. References
Bhatia, C.R. 1998. Terminator transgenics. Current Science 75:1288-1289.
Burk, D.L. 2004. DNA rules: legal and conceptual implications of biological lock-out
systems. California Law Review 92:1553-1587.
Daniell, H. 2002. Molecular strategies for gene containment in transgenic crops. Nature
Biotechnology 20:581-586.
Eaton, D., Van Tongeren, F., Louwaars, N., Visser, B. and Van der Meer, I. 2002.
Economic and policy aspects of 'terminator' technology. Biotechnology and Development
Monitor 49:19-22.
Federation of German Scientists and EcoNexus. 2006. GURTs: no case for field trials.
Convention on Biological Diversity (CBD) COP8, Curitiba, Brazil, March 2006. 4 pp.
Ellstrand, N.C., Prentice, H.C. and Hancock, J.F. 1999. Gene flow and introgression from
domesticated plants into their wild relatives. Annual Review of Ecological Systems
30:539-563.
Giovannetti, M. 2003. The ecological risks of transgenic plants. Rivista di Biologia
96:207-223.
Goeschl, T. and Swanson, T. 2003. The development impact of genetic use restriction
technologies: a forecast based on the hybrid crop experience. Environment and
Development Economics 8:149-165.
Gupta, P.K. 1998. The terminator technology for seed production and protection: why
and how? Current Science 75:1319-1323.
Kaiser, J. 2000. USDA to commercialize terminator technology. Science 289:709-710.
Kirkpatrick, S. and Tarasuk, V. 2003. The relationship between low income and
household food expenditure patterns in Canada. Public Health Nutrition 6:589-597.
Lawrie, R.G., Matus-Cadiz, M.A. and Hucl, P. 2006. Estimating out-crossing rates in
spring wheat cultivars using the contact method. Crop Science 46:247-249.
Lehmann, V. 1998. Patent on seed sterility threatens seed saving. Biotechnology and
Development Monitor 35:6-8.
Lence, S.H., Hayes, D.J., McCunn, A., Smith, S. and Niebur, W.S. 2005. Welfare
impacts of intellectual property protection in the seed industry. American Journal of
Agricultural Economics 87:951-968.

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Mahajan, S.K. 1999. Terminator technology a weapon of biological warfare. Current

Science 76:617.
Masood, E. 1998. Monsanto set to back down over terminator gene? Nature 396:503.
Matus-Cadiz, M.A., Hucl, P., Horak, M.J. and Blomquist, L.K. 2004. Gene flow in wheat
at the field scale. Crop Science 44:718-727.
Niiler, E. 1999. Terminator technology temporarily terminated. Nature Biotechnology
17:1054.
Pendleton, C.N. 2004. The peculiar case of terminator technology: agricultural
biotechnology and intellectual property protection at the crossroads of the third green
revolution. Biotechnology Law Report 23:1-29.
Pilger, G. 2002. Terminator could eliminate GM volunteers. Canola Guide May 2002:1617.
Rakshit, S. 1998. Terminator technology: science and politics. Current Science 75:747749.
Schiemann, J., Stamp, P., Foueillassar, X., Atanassov, A., Renard, M., Pierre, J., Gleba,
Y., Bock, R., Curtis, M. and Gressel, J. 2005. Biological containment systems for
genetically modified plants. Proc. of 2nd International Conf. on Co-existence between GM
and non-GM based agricultural supply chains. Montpellier, France. Edited by A.
Messean. Agropolis Productions, Montpellier.
Service, R.F. 1998. Seed-sterilizing terminator technology sows discord. Science
282:850-851.
van Wijk, J. 1994. Hybrids, bred for superior yields of for control? Biotechnology and
Development Monitor 19:3-5.
Visser, B. 1998. Effects of biotechnology on agro-biodiversity. Biotechnology and
Development Monitor 35:2-7.
Visser, B., van der Meer, I., Louwaars, N., Beekwilder, J. and Eaton, D. 2001. The
impact of 'terminator' technology. Biotechnology and Development Monitor 48:9-12.
Vom Brocke, K., Presterl, T., Christinck, A., Weltzien, R. and Gieger, H.H. 2002.
Farmers' seed management practices open up new base populations for pearl millet
breeding in a semi-arid zone of India. Plant Breeding 121:36-42.
United States Patent and Trademark Office, March 3, 1998.
http://patft.uspto.gov/netacgi/nphParser?Sect1=PTO1&Sect2=HITOFF&d=PALL&p=1&u=/netahtml/srchnum.htm&r=1&
f=G&l=50&s1=5,723,765.WKU.&OS=PN/5,723,765&RS=PN/5,723,765
Yadav, O.P., Weltzien-Rattunde, E., Bidinger, F.R. and Mahalakshmi, V.
2000. Heterosis in landrace-based topcross hybrids of pearl millet across arid
environments. Euphytica 112:285-295.

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Appendices
Appendix 1: Abstract patent for Control of plant gene expression (From United States
Patent and Trademark Office 1998).
A method for making a genetically modified plant comprising regenerating a whole plant
from a plant cell that has been transfected with DNA sequences comprising a first gene
whose expression results in an altered plant phenotype linked to a transiently active
promoter, the gene and promoter being separated by a blocking sequence flanked on
either side by specific excision sequences, a second gene that encodes a recombinase
specific for the specific excision sequences linked to a repressible promoter, and a third
gene that encodes the repressor specific for the repressible promoter. Also a method for
making a genetically modified hybrid plant by hybridizing a first plant regenerated from
a plant cell that has been transfected with DNA sequences comprising a first gene whose
expression results in an altered plant phenotype linked to a transiently active promoter,
the gene and promoter being separated by a blocking sequence flanked on either side by
specific excision sequences to a second plant regenerated from a second plant cell that
has been transfected with DNA sequences comprising a second gene that encodes a
recombinase specific for the specific excision sequences linked to a promoter that is
active during seed germination, and growing a hybrid plant from the hybrid seed. Plant
cells, plant tissues, plant seed and whole plants containing the above DNA sequences are
also claimed.

Inventors: Oliver; Melvin John (Lubbock, TX); Quisenberry; Jerry Edwin (Idalou,
TX); Trolinder; Norma Lee Glover (Quanah, TX); Keim; Don Lee (Leland,
MS)
Assignee: Delta and Pine Land Co. (Scott, MS); The United States of America as
represented by the Secretary of (Washington, DC)
Appl. No.: 477559
Filed:
June 7, 1995

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Appendix 2: Figure showing genetic basis of terminator technology for pure line seed
production in self-pollinated crops (From Gupta 1998).

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