EJB Electronic Journal of Biotechnology ISSN: 0717-3458 Vol.1 No.3, Issue of August 15, 1998.

1998 by Universidad Catlica de Valparaso - Chile Invited review paper/ Received 26 October, 1998

REVIEW ARTICLE

Critical role of plant biotechnology for the genetic improvement of food

crops: perspectives for the next millennium
Rodomiro Ortiz
The Royal Veterinary and Agricultural University (KVL), Department of Agricultural Sciences, Plant Breeding and Crop Science Section, 40
Thorvaldsensvej, DK-1871 Frederiksberg C, Copenhagen, Denmark.
Tel: +45 3528 3465 Fax: +45 3528 3468
E-mail: [email protected]

http:// www.agsci.kvl.dk/breed

This article reviews some of the highlights of modern forces, user demands, and public views cannot be ignored
plant biotechnology and discusses the potential when addressing basic and strategic research issues because
applications of biotechnology in the betterment of these factors shape scientific investigations and technology
farming systems in the next millennium. Plant or product development.
biotechnology will facilitate the farming of crops with
multiple durable resistance to pests and diseases, In writing this article the editor requested that I reflect on
particularly in the absence of pesticides. Likewise, the critical role that plant biotechnology may have in
transgenes or marker-assisted selection may assist in the assisting the genetic improvement of crops in the next
development of high yielding crops, which will be millennium. Within this context, I will discuss somewhat
needed to feed the world and save land for the philosophically, how biotechnology could help in solving
conservation of plant biodiversity in natural habitats. the increasingly enormous challenge of our time:
Hence, crops should be engineered to meet the demands adequately and appropriately feeding the world in a
and needs of consumers. The genetic base of crop sustainable manner.
production can be preserved and widen by an
integration of biotechnology tools in conventional This article restricts its discussion to gene-biotechnology,
breeding. Similarly targeting specific genotypes to mostly developed in the past 20 years, and not other
particular cropping systems may be facilitated by applications of non-gene biotechnology, which are known
understanding specific gene-by-environment to humankind for many hundred years ago. In addition I
interaction(s) with the aid of molecular research. High prefer to predict the potential applications of
quality crops with improved nutritional and health biotechnology in the genetic enhancement of crops only
characteristics as well as other aspects of added-value within the period of the coming decade. It would be
may be obtained through multidisciplinary co-operation inappropriate to attempt to provide an outlook beyond this
among plant breeders, biotechnologists, and other plant time-span because of the ever accelerating progress in this
scientists. Co-ordinated efforts between consumers, field. For example 15 years ago, plant biotechnology
policy makers, farmers and researchers will be required comprised only a few applications of tissue culture,
to convert the various aspects of a crop ideotype into recombinant DNA technology and monoclonal antibodies.
components of new and improved farming systems of Today, transformation, and marker-aided selection and
the next millennium. breeding are just a few of the examples of the applications
of biotechnology in crop improvement. This article was
written through the eyes of a classical geneticist (having
The end of a year, decade, century, or, as now, of a worked on the transmission of characteristics for the past 15
millennium, always offers an opportunity to reflect on years), and the practical view of a conventional plant
human activity in a particular discipline and to formulate a breeder, who has the desire to learn and accept innovative
future strategy. Researchers constantly examine past methods that enhance the available crop improvement
occurrences in order to learn lessons that could help in the techniques.
acquisition of new knowledge or for the further
development of appropriate technology ensuing from it. Of Background information
course, science and technology are not isolated in the
world, so researchers are expected to act according to the Writing about biotechnology for crop improvement in the
changing global society in which they live. This behaviour next millennium does not appear to be an easy task owing
could be seen as the major challenge of crop biotechnology to the rapid progress in this field. Within the last 100 years
for the next millennium, i.e., to consider the social actors in the world has seen the rise of genetics as a scientific
the research agenda and work. In other words, market discipline (1900s), the finding of DNA as the hereditary

This paper is available on line at http://www.ejb.org

Ortiz, R.

material (1944), the elucidation of the double helix 1998), where this economic activity still provides 60 to
structure of the DNA molecule (1953), the cracking of the 80% employment and 50% of national income (Anderson
genetic code (1966), the ability to isolate genes (1973), and 1996a). Such support will assist the developing world
the application of DNA recombinant techniques (from 1980 towards food self-reliance (Herdt 1998), which will be very
onwards). important to avoid hunger and keep peace in many regions
of the tropics, where the agricultural sector remains the
Methods of crop improvement have also changed most important basis for economic growth. Furthermore, a
dramatically throughout this century. Mass and pure line wealthy society provides high living standards to its
selection in landraces, consisting of genotype mixtures, citizens.
were the popular breeding techniques until the 1930s for
most crops. In the 1930s maize breeders started the Tissue culture was developed in the 1950s and became
commercial development of double cross hybrids that was popular in the 1960s. Today, micropropagation and in vitro
followed by the extensive utilization of single crop hybrids conservation are standard techniques in most important
since the 1960s (Troyer 1996). Pedigree-, bulk-, backcross- crops, especially those with vegetative propagation. At the
and other selection methods were also developed especially beginning of the 1980s genetic engineering of plants
for self-pollinating crop species. Such scientific advances in remained a promise of the future, although gene transfer
plant breeding led to the so-called Green Revolution, one had already been achieved earlier in a bacterium. The first
of the greatest achievements to feed the world in the years transgenic plant, a tobacco accession resistant to an
of the Cold War (Perkins 1997). Owing to this agricultural antibiotic, was reported in 1983. Transgenic crops with
betterment, cereal production, which accounts for more herbicide, virus or insect resistance, delayed fruit ripening,
than 50% of the total energy intake of the worlds poor, male sterility, and new chemical composition have been
kept in pace with the high average population growth rate released to the market in this decade (NCGR 1998; USDA-
of 1.8% since 1950 (Daily et al. 1998). Today, 370 kg of APHIS 1997). In 1996, there were about 3 million ha of
cereals per person are harvested as compared to only 275 transgenic crops grown in the world (mainly in North
kg in the 1950s; i.e., in excess of 33% per capita gain. America) whereas in excess of 34 million ha (a 12-fold
Similar progress in other food crops resulted in 20% per addition) of transgenic crops will be harvested this year in
capita gains since the early 1960s, according to FAO North America, Argentina, China, and South Africa among
(1995). There are 150 million fewer hungry people in the other countries. Argentina is the leading developing country
world today than 40 years ago, though there are twice as with an excess of 4 million ha of transgenic herbicide-
many human beings. Despite this splendid progress in crop resistant soybean. There are 4.4 million ha of transgenic
productivity, even greater progress must be made in order corn (14% of total acreage), 5 million ha of transgenic
to feed an additional two billion people by the early part of soybean (20%), and 1.6 million ha of transgenic canola
the 21st century (Anderson 1996a). Around 800 million (42%) grown only in North America (Moore 1998). It has
people are hungry today and another 185 million pre-school been calculated that in 1998 US farmers are growing over
children are still malnourished owing to lack of food and 50% of their cotton fields with transgenic seeds, the largest
water, or disease (Herdt 1998). Hence as suggested by the percentage for any crop ever. Trees are the next target in
Nobel Peace Laureate, Norman Bourlag (1997), new the agenda of genetic engineering.
biotechniques, in addition to conventional plant breeding,
are needed to boost yields of the crops that feed the world. Allozymes were available as the first biochemical genetic
markers in the 1960s. Population geneticists took advantage
Careful choice of such biotechniques as well as a realistic of such marker system for their early research. In the 1970s,
assessment of their potential in crop improvement are restriction fragment length polymorphisms (RFLP) and
needed to avoid not only the criticism of the anti-science Southern blotting were added to the tool box of the
lobbyists but also the permanent distrust of pragmatic geneticists. Taq polymerase was found in the 1980s, and the
traditional breeders (Simmonds 1997). For example, a polymerase chain reaction (PCR) developed shortly
World Bank panel recently released for discussion a well afterwards. Since then, marker-aided analysis based on
based report concerning bioengineering of crops (Kendall et PCR have become routine in plant genetic research and
al. 1997). In this working paper, the panel members marker systems have shown their potential in plant
recommend to give priority to all aspects of increasing breeding (Paterson 1996). Furthermore, new single
agricultural productivity in the developing world while nucleotide polymorphic markers based on high density
encouraging the necessary transition to sustainable DNA arrays, a technique known as gene chips (Chee et al.
methods. Indeed, plant biotechnology has been regarded as 1996), have recently been developed. With gene chips,
a priority area for technology transfer (Altman and DNA belonging to thousand of genes can be arranged in
Watanabe 1995), because genetically modified food, feed, small matrices (or chips) and probed with labeled cDNA
and fibre are of vital concern to the developing world (Ives from a tissue of choice. DNA chip technology uses
and Bedford 1998). Therefore, the rich industrialized world microscopic arrays (or micro-arrays) of molecules
should share their biotechniques and avoid policies that do immobilized on solid surfaces for biochemical analysis
not allow the progress of agriculture in poor, non- (Lemieux et al. 1998; Marshall and Hodgson 1998; Ramsay
industrialized parts of the world (Erbisch and Maredia 1998). An electronic device connected to a computer may

153
 Critical role of plant biotechnology for the genetic improvement of food crops: perspectives for the next millennium

read this information, which will facilitate marker-assisted The need for profit, as in any other business, has attracted
selection in crop breeding. In summary, since Mendels the interest of the private sector to defend their investments
work on peas, there have been five eras in genetic marker in crop biotechnology with patents, intellectual property
evolution (Liu 1997): morphology and cytology in early rights, and new protection methods, e.g. terminator
genetics (until late 1950s), protein and allozyme technology that inhibits germination of self-pollinated
electrophoresis in the pre-recombinant DNA time (1960 - seeds. This technology protection system prevents farmers
mid1970s), RFLP and minisatellites in the pre-PCR age from saving seeds from their harvest for further utilization
(mid 1970s - 1985), random amplified polymorphic DNA, as next season planting propagules. Three genes, each with
microsatellites, expressed sequence tags, sequence tagged a specific promoter, are inserted into the terminator plant
sites, and amplified fragment length polymorphism in the (D.E. Culley, Washington State Univ. in RAFI 1998). One
oligoscene period (1986 - 1995), and complete DNA of the genes (e.g. CRE/LOX system from bacteriophages)
sequences with known or unknown function as well as produces a recombinase that removes a spacer between the
complete protein catalogs in the current computer robotic gene producing, for example, a ribosomal inhibitor protein
cyber genetics generation (1996 onwards) The driving and its promoter such as late embryonic abundance, which
force for such a development has been the scientific interest only becomes active during the late stages of embryo
of human beings to understand and manipulate the development. This spacer with specific recognition sites
inheritance of their own characters. blocks the gene (for the ribosomal inhibitor protein) from
being activated. Another gene (e.g. tetracycline repressor
Responses to biotechnology in crop improvement system) produces a repressor that keeps off the recombinase
gene until an outside stimulus is applied to the terminator
The advances in plant transgenics and genomics described plant, e.g. a chemical such as the tetracycline, or
above have not been isolated from society (Busch et al. temperature and osmotic shocks. The United States
1991). Some of these achievements have been acclaimed by Department of Agriculture (USDA) and a cotton seed
end-users whereas other accomplishments, e.g. release of enterprise jointly acquired a patent for this concept (U.S.
genetically modified organisms (GMO), are being attacked, patent 5,723,765). Two months after this patent was
not only in words but also in deeds, by political activists. announced, one of the leading agro-chemical transnationals
Some of these educated middle-class campaigners are bought the cotton seed company, although one of its
expressing in this way their rampant eco-paranoia, while officers said that it may take many years before this
others hide their real agenda to manipulate the fashionable terminator gene idea becomes a proven technology in the
ecological movement. This controversy has attracted the seed industry.
attention of non-scientific partizans to each side. There
have been negative comments about transgenic plants by a Strategic alliances, joint ventures, research partnerships,
crown prince and contrasting positive comments by a new investments, company mergers, cross-ownerships, and
former president, both of whom may not have the required take-overs in the seed and agro-chemical business have also
technical knowledge to assess the potential of been in the news in recent months. Likewise, some leading
biotechnology for crop improvement. Irrespective of this scientists are leaving their academic appointments to join
ideological dispute and ensuing democratic disagreements, the new private enterprises in plant biotechnology. These
biotechnology products will be accepted by people who events are happening because the private sector wants to
support scientific-based progress, in a similar way that new use biotechnology to accelerate its growth in agri-business
cultivars or innovative crop husbandry techniques have in the short-term. Nonetheless, funds to support basic and
previously become integral parts of farming systems strategic research by public researchers are needed for a
elsewhere. However, without end-users consent, the long-term sustainable transfer of public goods (both
impact of a new technology in the society will be small or knowledge and technology) to the private sector or other
nil. users.

Scientific honesty seems to the best policy to convince Bioinformatics

people about the advantages of biotechnology for crop
improvement (Frewer et al. 1998). What to do? Scientists, Another important factor in the successes of the genetic
farmers, consumers, and policy-makers should objectively improvement of crops was the development of fast and
assess the potential hazards of crop biotechnology in more reliable computers, which allowed easier management
farming and food systems regarding the current situation and analysis of data as well as publication of scientific
and the likelihood that such hazards may occur. For reports. The impact of the informatic revolution in crop
example scientists should explain to the people that gene improvement can be partially assessed by counting the
recombination (or reassortment) already occurs in nature. number of publications indexed in Plant Breeding Abstracts
However, the ecological success of viable recombinants (CAB International, Wallingford, Oxon, UK). There was
after gene reassortment is unpredictable owing to the high ca. 22-fold increase of publications in the 1930-1997 period
fitness of current isolates. For this reason, more scientific (Fig. 1). It was in the 1970s that indexed publications in
research will be needed to identify unpredictable risks and plant breeding exceeded 10,000 per year. More publications
the chances of their occurrence. and easy means for retrieving this information accounted

154
Ortiz, R.

for such growth of knowledge dissemination in plant Molecular biologists will study not only individual genes
genetics and breeding. Today, rapid information exchange but how circuits of interacting genes in different pathways
has been facilitated with electronic mail and access to the control the spectrum of genetic diversity in any crop
internet to read electronic publications such as this journal. species. For example, more information will be available on
Nowadays, information technology and DNA science are why plant resistance genes are clustered together, or what
beginning to fuse into a single operation. Computers are candidate genes should be considered when manipulating
deciphering, and organizing the huge genetic information quantitative trait loci (QTL) for crop improvement
that may become the raw resource of the emerging biotech (Paterson 1997).
economy in the next century (Rifkin 1998). Scientists
working in the new field of bioinformatics are developing Farming in environmentally friendly systems
biological data banks to download the genetic information
accumulated during millions of years of life evolution, and The aims of applied plant science research for agriculture
perhaps reconstruct some of the living organisms of the are to enhance crop yields, improve food quality, and
natural world. preserve the environment where human beings and other
organisms live. The best way for conservation of plant
biodiversity and its environment, would be to achieve high
crop productivity per unit area. In this regard, Briggs (1998)
reported that as yields treble, soil erosion per ton of food
decreases by two-thirds. There has been a significant yield
improvement owing to enhanced crop husbandry, but in the
next years progress will be achieved by changing plants that
could be more suitable to sustainable and environmentally
friendly farming systems. Agro-chemical corporations are
developing pest and disease resistant transgenic crops to
avoid pollution with pesticides in the farming system.
Furthermore, food quality will become more important than
crop productivity in a wealthy society. Consumers will
prefer transgenic crops if they have the desired
characteristics.
Figure 1. Number of publications indexed in Plant
Breeding Abstracts (CAB International, Wallingford, Oxon, In the next decades meiotic-based breeding will still
UK) since its publication in early 1930s generate cultivars for farmers. Genetic improvement
through biotechnology needs conventional breeding
because (1) the elite cultivars will be the parents of the next
Plant genomics generation of improved genotypes, (2) field testing across
locations or cropping systems and over years will be needed
This new term, defined by the development of to determine the best selections due to the genotype-by-
biotechnology, refers to the investigations of whole environment interaction (Kang and Gauch 1996). As stated
genomes by integrating genetics with informatics and by Briggs (1998), transgenes must be viewed as
automated systems. Genomic research aims to elucidate the improvements rather than replacements for elite
structure, function and evolution of past and present germplasm. Indeed, genetic engineering may provide a
genomes (Liu 1997). Some of the most dynamic fields means to add value by introducing synthetic or natural
concerning agriculture are the sequencing of plant genes that enhance crop quality and yield, as well as protect
genomes, comparative mapping across species with genetic the plant against pest and diseases. Farmers will pay more
markers, and objective assisted breeding after identifying for transgenic crop propagules if they obtain extra-income
candidate genes or chromosome regions for further after adopting biotech-derived products. For example, seeds
manipulations. As a result of genomics, the concept of gene of insect resistant transgenic crops will be more expensive
pools has been enlarged to include transgenes and native than those of available cultivars but the farmer will not
exotic gene pools that are becoming available through need to apply pesticides in their transgenic fields. Of
comparative analysis of plant biological repertoires (Lee course, patents make transgenic seeds more expensive but
1998). Understanding the biological traits of one species also farmers benefits may be higher.
may enhance the ability to achieve high productivity or
better product quality in another organism. Gene banks, DNA banking and virtual plant
breeding
DNA markers and gene sequencing provides quantitative
means to determine the extent of genetic diversity and to The sequencing of crop genomes opened new frontiers in
establish objective phylogenetic relationships among conservation of plant biodiversity and its genetic
organisms. Gene chips and transposon tagging will enhancement. The advances in gene isolation and
provide new dimensions for investigating gene expression. sequencing in many plant species allows to envisage that

155
 Critical role of plant biotechnology for the genetic improvement of food crops: perspectives for the next millennium

within a few years, gene-bank curators may replace their of first mapping a gene to subsequently determine its exact
large cold stores of seeds with crop DNA sequences that function. Gene identification in DNA libraries coupled with
will be electronically stored. The characterization of plant biosource technology and an enhanced ability to put genes
genomes will ultimately create a true gene bank, which into plants will be routine for improving crops in the next
should possess a large and accessible gene inventory of decade.
todays non-characterized crop gene pools. Of course, seed
banks of comprehensively investigated stocks should Genomics may provide a means for the elucidation of
remain because geneticists and plant breeders, the main important functions that are essential for crop adaptedness
users of gene banks, will need this germplasm for their (Wallace and Yan 1998). Regions of the world should be
work. Genomics may accelerate the utilization of candidate mapped by combining data of geographical information
genes available at these gene banks through transformation systems, crop performance, and genome characterization in
without barriers across plant species or other living each environment. In this way, plant breeders can develop
kingdoms. Nonetheless, genetic engineering should be seen new cultivars with the appropriate genes that improve
as one of the methods of plant breeding that permits the fitness of the promising selections. Fine-tuning plant
direct alteration and re-building of a crop population. responses to distinct environments may enhance crop
Shutting-off genes coding for undesired characteristics productivity. Development of cultivars with a wide range of
may be another application of transgenics in crop adaptation will allow farming in marginal lands. Likewise,
improvement. research advances in gene regulation, especially those
processses concerning plant development patterns, will help
Plant breeders will change their modus operandi with the breeders to fit genotypes in specific environments.
development of objective marker-assisted introgression and Photoperiod insensitivity, flowering initiation,
selection methods. Backcross breeding will be shortened by vernalization, cold acclimation, heat tolerance, host
eliminating undesired chromosome segments (also known response to parasites and predators, are some of the
as linkage drags) of the donor parent or selecting for more characteristics in which advanced knowledge may be
chromosome regions of the recurrent parent. Parents of elite acquired by combining molecular biology, plant physiology
crosses may be chosen based on a combination of DNA and anatomy, crop protection, and genomics.
markers and phenotypic assessment in a selection index, Multidisciplinary co-operation among researchers will
such as best linear unbiased predictors (Bernardo 1998). To provide the required holistic approach to facilitate research
achieve success in these endeavours, cheap, easy, progress in these subjects.
decentralized, and rapid diagnostic marker procedures are
required. Pharming and Farmer-ceuticals

There are many areas of basic and strategic research in Growth of cities in the developed world has already
plant breeding and genetics that are being facilitated by replaced farmland with shopping malls, parking lots, and
marker-aided analysis (Paterson 1996). With molecular housing developments. Peri-urban agriculture and home
markers, plant biologists are reviewing crop evolution and gardening are also becoming very important for national
gathering new knowledge. Such information should be food security in the developing world as a result of rapid
incorporated into genetic enhancement programmes, urban expansion. Hence, new cultivars will be needed to fit
especially those with an evolutionary breeding scheme. into intensive production systems, which may provide the
Likewise, plant ideotypes for each crop should drive the food required to satisfy urban world demands of the next
work of plant breeders. Specific plant morphotypes have century. Specific plant architecture, tolerance to urban
been defined in rice and wheat based on accumulated pollution, efficient nutrient uptake, and crop acclimatization
knowledge of crop physiology and crop protection. The to new substrates for growing are, among others, the plant
needed characteristics required to develop improved plant characteristics required for this kind of agriculture. Genes
prototypes ensuing from such a virtual breeding approach controlling these characteristics may be available in gene
may be available in gene banks of the crop or in those of banks for further cross breeding, which can be assisted by
other species. Otherwise, breeders may obtain novel genomics. Peri-urban and home garden farmers will have
transgenes to develop the required ideotype. to adapt to new demands from emerging urban populations
with higher income. These consumers may request a more
Nowadays, the finding of new genes that add value to varied diet. For example, food crops with low fats, and high
agricultural products seems to be very important in the in specific amino acids may be needed to satisfy people
private agri-business. Unique gene databases are being who wish to change their eating habits. If genes controlling
assembled by the industry with the massive amount of data these characteristics do not exist in a specific crop pool they
generated by genomics research. A new term biosource may be incorporated into the breeding pool using
was coined recently to refer to a fast and effective licensed transgenics.
technology of pinpointing genes. With this method, a
benign virus infects a plant with a specific gene that Some publications anticipated that in the next millennium
allows researchers to observe directly its phenotype. food will not need to be harvested from farmers fields
Biosource replaces the standard time-consuming approach (Anderson 1996b). Tissue culture of certain parts of the

156
Ortiz, R.

plant may provide a means to achieve success in this viability. People also want that their views about
endeavour. For example, edible portions of fruit crops applications of biotechnology for improving agriculture are
could be grown in vitro. A steady and cheap supply of these listened irrespective of their knowledge in the subject.
edible plant parts will be required in this new agri-business. Moreover, farmers are afraid that negative propaganda
It will take some time before such a process can be scaled- jeopardizes the public image of their products. Scientists
up for commercial output. Nonetheless, a patent was and policy markers should not forget that peoples
submitted in 1991 by a Californian biotech company for acceptability is the most important component of the
producing a vanilla extract through cell culture. Of course, general public assessment of risk, which includes both
this technique will not replace farming as we know it today. uncertainty and negative consequences. This acceptability
This biotechnique, as well as other new farming methods, depends on cultural factors because peoples views change
offers a means for new ways of producing food, feed or according to time and location.
fibre.
The process of risk assessment in agro-chemical consists of
Often plants provide the raw materials for agro-industry, (i) hazard identification, (ii) exposure assessment, (iii)
and not only for food or fibre processing. Active effects management, (iv) risk characterization, and (v) risk
ingredients of plants have been transformed into management. However, transgenic crops may be able to
commercial products such as medicines, solvents, dyes, and invade (or colonize) and multiply in many habitats. Hence,
non-cooking oils for many years. Hence, it would not be this risk assessment of a genetically modified living
surprising to see, in few years from now, entire farms organism (also known as GMLO) must consider other
without food crops but growing transgenic plants to characteristics not included when assessing the release of
produce new products, e.g. edible plastic from peas or plant non-living compounds to the environment, e.g. horizontal
oils to manufacture hydraulic fluids and nylon (Grace gene transfer between transgenic crops and wild related
1997). This new rural activity may result in important species. Scientific risk assessment of transgenic crops must
changes in the national economic sector. be strictly performed and precautionary principles should
be considered in the decision making process. In the
Pharming has been added to the dictionary to indicate a industrialized world, this precautionary principle is a key
new kind of system to obtain medicines (Anderson 1996b). component of the response to the unforeseen (and
For example, oral vaccines appear to be a convenient sometimes irreversible) human and environmental impact,
delivery system for vaccination throughout the world. which may occur by introducing into the system new
Biotechnology has been used to engineer plants that contain advances ensuing from research and technology
a gene derived from a human pathogen (Tacker et al. 1998). development. In Norway, an unique legislation advocates
An antigenic protein encoded by this foreign DNA can that the production and use of GMO should be ethically
accumulate in the resultant plant tissues. Results from pre- and socially justifiable in accordance with the principle of
clinical trials showed that antigenic proteins harvested from sustainable development as well as safe to humans and to
transgenic plants were able to keep the immunogenic the environment. By applying this framework, marketing
properties if purified. These antigenic proteins caused the applications of GMO could be rejected if insufficient
production of specific antibodies in injected mice. Mice, documentation regarding ecological and heath aspects was
which ate these transgenic plant tissues, also showed also a submitted by the producer.
mucosal immune response. Arakawa et al. (1998) recently
demonstrated the ability of transgenic food crops to induce What are the potential ecological risks associated with the
protective immunity in mice against a bacterial enterotoxin release of GMP into the farming system? These are of
such as cholera toxin B subunit pentamer with affinity for course a very large number of potential risks, However,
GMI-ganglioside. Also, potato tubers have been used perhaps the two most important risks are:
successfully as a biofactory for high-level output of a (i) GMP establishes in semi- or natural habitats, and
recombinant single chain antibody (Artsaenko et al. 1998). (ii) inserted transgenes incorporate into other species,
thereby affecting non-target organisms in farms or
Risk assessment of transgenic crops natural habitats.
Hierarchical test protocols have been proposed to assess the
Lack of scientific data, non-scientific partizan views, risks of releasing GMP. Such protocols require knowledge
uncertainty of potential risks, and ignorance confound about evolutionary history, morphology, life-history
rational discussion concerning the release of GMO. The characteristics, pollination or breeding system, gene-
issue of releasing genetically modified plants (GMP) into transfer likelihood, natural hybridization, recruitment and
the farming system has become particularly agitated by vegetative propagation of a chosen species. Likewise,
lobbyist groups in Europe despite widespread cultivation of producers should provide, to facilitate this risk assessment,
such crops in North America and elsewhere. Scientists must additional information regarding biochemical,
realise that the general public are concerned that an physiological, and morphological changes owing to inserted
uncautious approach to the manipulation and cultivation of gene(s), along with a list and description of marker and
transgenic crops may affect biodiversity and its sustainable reporter genes included in the transgenic plant. It would
utilization in the farming system, e.g. loss of variability and also be important to add details concerning when and in

157
 Critical role of plant biotechnology for the genetic improvement of food crops: perspectives for the next millennium

which plant tissues or organs will be expressed the agriculture will be easily resolved in a democratic society.
modified function or phenotype. Nonetheless, people must The public need to choose between being safely self-
also know that scientists assessing risks of transgenic crops regulated or to follow safety regulations as agreed by
may extrapolate the outcome or results from simple short- lawmakers after listening to the views of scientists,
time experiments into complex long-term natural- or producers, and consumers.
farming systems. Investigations about gene flow and The general public should see biotechnology as a safe tool
competing ability of transgenic crops may be easily for scientific crop improvement, because it helps in the
addressed through short-term experiments. However, the fight against hunger and poverty. Therefore, research
assessment of the environmental impact of GMP requires a funding should be allocated accordingly to long-term plant
long-term, expensive, holistic research. Computer breeding programmes, which include biotechnology as one
modelling, which integrates knowledge about gene flow, of its tools. In this way, we may effectively face the serious
competing ability, spread of transgenes to weedy species, challenge of feeding the rapidly growing world population
and cultural practices in the farming system, may provide in the next millennium.
an alternative means for long-term risk assessment of
releasing GMP into the environment. Acknowledgments
Consumer concern about transgenic crops also focuses on
their safety as food, especially if modifications could Thanks to the Nordic Council of Ministers for providing
influence their metabolism or health. In this regard, funding support to the author via a Nordic Professorship in
transgenic plants without selectable markers, such as Plant Genetic Resources, and to Drs. Jonathan H. Crouch
antibiotic resistance genes, are needed to convince GMP- (Elsoms Seeds Ltd., Spalding, Lincolnshire, England) and
sceptics of the advantages of genetic engineering for crop Lise Nrrind Hansen (formerly at KVL, Denmark) for
improvement. In this way, their criticism concerning the comments during the development of this paper.
potential risks of transgenic crops could be overcome. For
example, molecular or metabolic markers may provide a References
means to identify transgenic plants with desired trait(s). Of
course, these alternative markers should be safe from an Altman, D.W. and Watanabe, K.N. (1995).Plant
environmental and health perspective. biotechnology transfer to developing countries. Academic
Press, Inc., California and London.
Outlook
Anderson, J. (1996a). Feeding a hungrier world.
Within the next 10 or 20 years, five research areas may Phytopathology News 30 6: 90-91.
become very important for crop improvement: (i) apomixis
to fix hybrid vigour, (ii) male sterility systems with Anderson, W.T. (1996b). Evolution isnt what is used to be.
transgenics for hybrid seed in self-pollinating crops, (iii) W.H. Freeman and Company, New York. 223 pp.
parthenocarpy for seedless vegetables and fruit trees, (iv)
short-cycling for rapid improvement of forest and fruit Arakawa, T., Chong, K.X. and Langridge, W.H.R. (1998).
trees, and (v) converting annual into perennial crops for Efficacy of a food plant-based oral cholera toxin B subunit
sustainable agricultural systems. The development of vaccine. Nature Biotechnology 16: 292-297.
perennial crops will be especially important to protect the
soil from erosion. Plant biotechnology will play, of course, Artsaenko, O., Kettig, B., Fiedler, U., Conrad, U. and
an important role in achieving research and development Dring, K. (1998). Potato tubers as a biofactory for
success in these areas. recombinant antibodies. Molecular Breeding 4: 313-319.
Banning transgenic crops in the farming system will be
foolish because the potential benefits are so great. Bernardo, R. (1998). A model for marker-assisted selection
Environmentalists should recall or re-read Silent Spring among single crosses with multiple genetic markers.
by Rachel Carlson (1962). Whatever scientists do to Theoretical Applied Genetics 97: 473-478.
develop crops that eliminate or reduce the utilization of
polluting agro-chemicals in the farming systems must be Bourlag, N.E. (1997). Feeding a world of 10 billion people:
welcome by farmers and consumers. For example, one the miracle ahead. Plant Tissue Culture and Biotechnology
interesting approach for developing resistant transgenic 3: 119-127.
crops may be through the improvement of the plants own
defence system. Inducible and tissue specific promoters Briggs, S.P. (1998). Plant genomics: more than food for
could assist in this endeavour. thought. Proceedings of the National Academy of Sciences
Collective approval may lead to new partnerships, co- USA 95: 1986-1988.
operation or joint ventures in research and development
between scientists in the public and private sectors that will Busch, L., Lacy, W.B., Burkhardt, J. and Lacy R.L. (1991).
benefit farmers and consumers with profits and high quality Plants, power, and profit: social, economic, and ethical
products, respectively. Any potential risk in human consequences of the new biotechnologies. Basil Blackwell
development associated with biotechnology applications in Inc., Cambridge, Massachussetts. 275 pp.

158
Ortiz, R.

Carlson, R. (1962). Silent spring. Houghton Mifflin Co.,

Boston. Liu, B.L. (1997). Statistical genomics: linkage, mapping,
and QTL analysis. CRC Press, Boca Raton. 611 pp.
Chee, M., Yang, R., Hubbell, E., Berno, A., Huang, X.C.,
Stern, D., Winkler, J., Lockhart, D.J., Morris, M.S. and Marshall, A. and Hodgson, J. (1998). DNA chips: an array
Fodor, S.P.A. (1996). Accessing genetic information with of possibilities. Nature biotechnology 16:27-31.
high-density DNA arrays. Science 274: 610-614.
Moore, S.D. (1998). Agrochemical rivalry heats up: AHP-
Daily, G., Dasgupta, P., Bolin, B., Crosson, P., du Guerny, Monsanto pact raises pressure. The Wall Street Journal
P., Ehrlich, P., Folke, C., Jansson, A.M., Jansson, B.-O., Europe, 5-6 June 1998, p. 10
Kautsky, N., Kinzig, A., Levin, S., Mler, K.G., Pinstrup-
Andersen, P., Siniscalco, D. and Walker, B. (1998). Global NCGR. (1998).
food supply: food production, population growth, and the http://www.ncgr.org/gpi/oddysey/agbio/foods.html
environment. Science 281: 1291-1292.
Paterson, A.H. (ed.) (1996). Genome mapping in plants.
Erbisch, F.H. and Maredia, K.M. (eds.) (1998). Intellectual Academic Press, Inc. and R.G. Landes Co., New York and
property rights in agricultural biotechnology. Austin. 330 pp.
Biotechnology in Agriculture 20. CAB International,
Wallingford. 240 pp. Paterson, A.H. (ed.) (1997) Molecular dissection of
complex traits. CRC Press, Inc., Boca Raton and New
FAO (Food and Agricultural Organization of the United York. 305 pp.
Nations). (1995). Dimensions of need: an atlas of food and
agriculture. FAO, Rome, Italy. Perkins, J.H. (1997). Geopolitics and the green revolution.
Oxford University Press, New York. 337 pp.
Frewer L.J., Howard C. and Shepherd, R. (1998). The
influence of initial attitudes on responses to communication RAFI (Rural Advancement Foundation International)
about genetic engineering in food production. Agriculture (1998) RAFI Communique March/April 1998: How does
and Human Values 15: 15-30. the terminator technology work? pp. 4-5.

Grace, E.S. (1997). Biotechnology unzipped: promises & Ramsay, G. (1998). DNA chips: state of the art. Nature
realities. Joseph Henry Press, Washington D.C. 248 pp. Biotechnology 16:40-44.

Herdt, R.W. (1998). Assisting developing countries toward Rifkin, J. (1998). The biotech century. Victor Gollancz,
food self-reliance. Proceedings of the National Academy of London. 272 pp.
Sciences USA 95: 1989-1992.
Simmonds, N.W. (1997). Pie in the sky. Tropical
Ives, C. and Bedford B. (eds.) (1998). Agricultural Agriculture Association June 1997: 1-5.
biotechnology in international development. Biotechnology
in Agriculture 21. CAB International, Wallingford. 368 pp. Tacker, C.O., Mason, H.S., Losonsky, G., Clements, J.D.,
Levine, M.M. and Arntzen, C.J. (1998). Immunogenecity in
Kang, M.S. and Gauch Jr., H.G. (eds.) (1996). Genotype- humans of a recombinant bacterial antigen delivered in a
by-environment interaction. CRC Press, Inc., Boca Raton. transgenic potato. Nature Medicine 4: 607-609.
416 pp.
Troyer, A.F. (1996). Breeding widely adapted, popular
Kendall, H.W., Beachy, R., Eisner, T., Gould, F., Herdt, R., maize hybrids. Euphytica 92: 163-174.
Raven, P.H., Schell, J.S. and Swaminathan, M.S. (1997).
Bioengineering of crops: report of the World Bank panel on USDA-APHIS (United States Department of Agriculture -
transgenic crops. Environmentally and Socially Sustainable Animal and Plant Health Inspection Service). (1997).
Development Studies and Monographs Series 23. The Biotechnology permits database.
World Bank, Washington D.C. 30 pp. http://www.aphis.usda.gov:80/bbep/bp/

Lee, M. (1998). Genome projects and gene pools: New Wallace, D.H. and Yan, W. (1998). Plant breeding and
germplasm for plant breeding? Proceedings of the National whole-system crop phyisiology: improving adaptation,
Academy of Sciences USA 95: 2001-2004. maturity, and yield. CAB International, Wallingford, Oxon.
390 pp.
Lemieux, B., Aharoni, A. and Schena, M. (1998). Overview
of DNA chip technology. Molecular Breeding 4: 277-289.

159