Induced mutations for enhancing nutrition and food

production

S. Mohan Jain* and P. Suprasanna**

*Department of Agricultural Sciences, University of Helsinki, PL-27, Helsinki, Finland

Email: mohan.jain@helsinki.fi

**Nuclear Agriculture & Biotechnology Division, Bhabha Atomic Research Centre,

Trombay, Mumbai 400 085. Email: prasanna@barc.gov.in

Received March,15 , 2011

Accepted May, 14, 2011

Geneconserve 40 : 201 – 215 (2011)

Abstract

Issues of enhanced food security depend primarily on increasing agricultural production.

Given this challenge, existing and new, appropriate technologies need to be integrated

into agricultural research, to focus on the problems related to improving nutritional

security. Among the different approaches, mutagenesis and the isolation of improved or

novel phenotypes in conjunction with conventional breeding programmes can result in

mutant varieties endowed with desirable traits. Induced mutations play an important role

enhancing nutritional quality in crop plants. Several mutant genes have been successfully

introduced into commercial crop varieties that significantly enhance the nutritional value

of crops. This review briefly outlines the aspects of induced mutations and nutritional

quality in crop improvement.

Introduction

Agricultural development has always been on the move towards increasing crop

productivity and exploiting natural resources. Such a developmental program necessitates

coordination between agricultural activities, ecosystems, and human society. It is

imperative that sustainable use of natural resources should be prudently managed in

conjunction with the advancement in the knowledge gained from science and technology.

Today human population is growing rapidly especially in the developing countries

whereas in the developed countries the situation is just the reverse. Global food security

continues to be the centre stage issue and plant breeders are under pressure to sustain the

food production to meet the demand of ever-growing human population. Further to the

problem, the erratic climatic change because of its direct effects on both food production

and food security has mounted the pressure to develop sustainable (Jain 2010a). Several

factors such as abiotic and biotic stresses, industrial pollution, deforestation, loss of

genetic diversity, water shortages and so on are responsible for having a negative impact

on food production. The world population is expected to reach 7 billion in the next 20

years, and 10 billion by 2050. The question is, Can we feed the world and sustain

Article

nutrition balance in the diet at the affordable price with the available technologies? The

answer seems to be not so positive even though plant breeders are making all out efforts

to sustain food production and nutrition to make genetic improvement of plants by using

conventional and modern tools. Pundit Jawaharlal Nehru, the great former prime minister

of India, remarked in the late 50s that “Everything else can wait but not agriculture” and

this remark remain relevant today. There is no short-term magic formula to solve the

world’s food problems.

The genetic variability is highly desirable for developing new cultivars, which is induced

by mutagen treatments and natural spontaneous changes. The spontaneous mutation rate

is pretty low and can’t be exploited for breeding and that is why artificially mutations are

induced with physical and chemical mutagen treatment. Quite many useful genetic

changes have been induced by mutagen treatment including high yield, flower colour,

disease resistance, and early maturation and so on in crop, vegetables, medicinal herbs,

fruit and ornamental plants. So far, over 3000 mutant varieties have been officially

released over 60 countries including rice, wheat, barley, sorghum, legumes, cotton, edible

oil, ornamental plants and fruits (www-mvd.iaea.org). China and India are the major

producers of mutant varieties to feed their ever-growing human population. Among all

crops, the released highest number of mutant varieties is in rice. In 2005, The

International Atomic Energy Agency (IAEA), Vienna, Austria was conferred Nobel Peace

Prize for its contributions to the peaceful applications of nuclear energy in various fields

including food and agriculture. This award was a shot in the arms by recognising the

major contributions made with the use of nuclear energy in enhancing food production

and economic benefits worldwide. The year 2008 marked the 80th

anniversary of mutation

induction in crop plants, when an international symposium on induced mutations in plants

was organised in Vienna, Austria (Shu, 2010).

The foremost objective of plant breeders and geneticists is to sustain food and nutrition

security and that is why the selection of major crops has become crucial for meeting these

goals under the existing arable land, and climate change. Malnutrition with respect to

micronutrients like vitamin A, iron, and zinc affects more than 40% of the world’s

population. Micronutrient deficiencies are common in many developing countries and are

typically due to inadequate food intake, poor dietary quality and poor bioavailability (Fig.

1, Ramakrishnan 2002). For example in wheat, zinc is quite low in grain and

consequently deficient in human diet among the developing countries, and to supplement

it needs zince enriched wheat grains at the farmer’s field (Hussain et al 2010). World

major crops like rice, wheat, maize, barley etc., would require continuous modifications

for sustainable food production, where as nutrition security would be maintained by

improving legumes, vegetables, and fruits. For example banana and plantain are among

the world’s major food crops, and are considered as the poor man’s fruit crop, and have

potential to provide subsistence diet and nutrition to millions of people. However, the

major problem with fruit breeding work is long life cycle of many fruit crops, which

varies from 3-25 years or even more. The large juvenile period has hampered fruit

breeding work. In fruit crops, mutagenesis has been quite useful in isolation of useful

mutants such as plant size, blooming time, fruit ripening, fruit colour, and resistance to

pathogens. Another major fruit crop is date palm (Phoenix dactylifera L.), a major source

of human nutrition including vitamins, sugars, fat, salts and minerals, and oils; and has

high potential to produce bio-ethanol (Jain 2011; Jain et al 2011).

Induced mutations

Mutations are induced by physical (gamma radiation, high and low energy beams) and

chemical (ethyl methane sulfonate, EMS) mutagen treatment of both seed and vegetative

propagated crops. The mechanism of mutation induction is that the mutagen treatment

breaks the nuclear DNA and during the process of DNA repair mechanism, new

mutations occur randomly and are heritable. The changes can also occur in cytoplasmic

organelles and also results in chlorophyll mutations, chromosomal or genomic mutations

that enable plant breeders to select useful mutants such as abiotic and biotic stresses and

others. By induced mutations, mutants with multiple traits can be identified. The chances

of survival varieties are much higher under the climate change. In Vietnam, eight rice

mutant varieties have been developed with multiple traits like high quality, tolerance to

salinity and short duration allowing three harvests per year providing farmers an extra

income of 300 million US dollars. Moreover, mutant varieties are readily accepted by the

consumers. Induced mutations have played a pivotal role in enhancing world food

security, since new food crop varieties with various induced mutations have contributed to

the significant increase of crop production at locations people could directly access

(Kharkwal and Shu 2010).

Among physical mutagens, gamma radiation has been widely used for mutation induction

for both seed and vegetative propagated crops. Recently ion energy technology- heavy ion

beam (HIB) and low energy ion beam (LIB)- is being for mutation induction in wide

ranging crops. HIB is predominantly used for inducing mutations in plants (Jain, 2010a).

They transfer linear energy transfer (LET) and enhances the induction of higher biological

effects. Several Arabidopsis mutants have been obtained including deletions, insertions,

and chromosomal translocations by HIB.

Plant cell tissue culture has made tremendous progress towards plant regeneration from

all major food and horticultural crops. Micropropagation via organogenesis is routinely

used for clonal propagation of ornamental plants and other vegetative propagated plants,

especially woody and fruits trees. Explant (shoot meristem, adventitious buds, and

microspores) is directly treated with mutagen and direct shoots are regenerated followed

by root formation (Suprasanna et al. 2010). Regenerated plants are maintained in the

greenhouse and put under the selection pressure. Similarly somatic embryogenesis of

vegetative propagated crops (banana, date palm, cassava and others) are readily induced.

The advantage of mutagen treatment to embryogenic cell suspension that chimeras are

either eliminated or dramatically reduced, and could get mutant somatic embryos which

are regenerated into plantlets. Embryogenic cells are plated on a filter paper and put on

agar solidified medium; treat cells with gamma radiation followed by transfer them on

culture medium and allow them to form somatic embryos. The treated cells can also be

put under the selection pressure in order to isolated mutants, e.g. disease, salt, drought

tolerant mutants. The selected mutant plants are transferred in the greenhouse and finally

to the field evaluation and use them for crossing with other varieties. The radiosensitive

curve should be determined to calculate LD50 dose (Lethal dose) for each experimental

plant to avoid either very high or very low dosage. Moreover, plants and even varieties

differ in radio sensitivity. Low dose of gamma radiation has promoted growth in citrus

depending of cultivar (Fig. 2), maintain embryogenic nature of date palm for 2-3 years,

promote growth in orchids, enhances secondary metabolites in medicinal plants, and used

for improving shelf life of post harvest products

A range of several mutants in different ornamental plants, maize, rice and wheat have

been isolated and used for crop breeding. Similarly LIB has been used for mutation

breeding and gene transfer. This method has many advantages such as low damage rate,

higher mutation rate and wider mutation spectrum. In rice, 11 new lines of rice mutants

with higher yield, broader disease resistance, and shorter growing period and high grain

quality were developed and now being cultivated in China. In jasmine rice from Thailand,

a wide range of mutants were recorded including short stature, red/purple colour of leaf

sheath, collar, auricles, ligules and dark brown stripes on leaf blade, dark brown seed coat

and pericarp.

Mutation induction for quality and nutrition improvement

Besides increase in yield of a crop, quality and nutrition components are equally

important in human diet. There is a necessity to enhance mineral elements (bio-

fortification) and amino acids essential for human and animals, alteration of protein and

fatty acids profiles for nutritional and health purposes, change of physicochemical

properties of starch for different end uses, enhancement of phyto-nutrients in fruits and

reduction of anti-nutrients in staple food. Induced mutations could play an important role

in inducing mutations for enhancing nutritional quality in crop plants. Of the 3000 mutant

varieties developed globally, 776 mutants (Fig. 3) have been induced for nutritional

quality (www-mvd.iaea.org).

Collaborative research programme under Food & Agriculture Organization and

International Atomic Energy Agency (FAO/IAEA) has been focussed on at crop

improvement by induced mutation using nuclear techniques (Jain 2000) intended to

produce strains of cereals with higher concentrations of micronutrients and improvement

of their bioavailability by reduction in the concentration of phytic acid. In this regard,

strategies should be aimed at breeding plants that can contain high levels of minerals and

vitamins in their edible parts to reduce substantially the recurrent costs associated with

fortification and supplementation (Shetty 2009). Such a strategy will be successful

depending on farmer’s willingness to adopt such varieties, palatability of the edible parts

of these varieties and consumer acceptability, and if the incorporated micronutrients can

be absorbed by the human body (Bouis 2002). Certain considerations need to be

addressed before a plant breeding strategy can be put in place to combat micronutrient

deficiency to function and to have universally adoptability, particularly in Developing

Countries (Bouis 2002). These include, feasibility to breed micronutrient- dense staple

food varieties, effects on plant yields and farmer’s adoption of such varieties, possibility

of changes that micronutrient density can have a great nutritional balance on the staple

diet of consumers, bioavailability of extra micronutrients in staple foods to humans, and

alternate options for more easily sustainable strategies for reducing micronutrient

malnutrition.

Several mutant genes have been successfully introduced into commercial crop varieties

that significantly enhance the nutritional value of crops like maize, barley, soybean, and

sunflower. In maize, quality protein maize (QPM) varieties are grown on hundreds of

hectare land. They have almost twice higher amount of lysine and tryptophan, and 30%

less leucine as compared to parental lines; shown a dramatic effect on human and animal

nutrition, growth and performance. In cassava, three mutants have been isolated showing

different size of starch grain. They have high economic potential for industrial use of

starch and influence on cooking quality. Small starch grain size seems to be highly

suitable for bio-ethanol production. In sweet sorghum, a mutant variety Yuantian No.1

has been developed in China (Fig. 4), which has 20% more total carbohydrates as

compared to the parental lines; well suited for Food, Feed and Bio-energy (three in one).

Five rice giant embryo mutants, characterized by enlarged embryo than that of wild type

were found to have increase in the contents of protein, vitamin B1, vitamin B2, vitamin E,

essential amino acids such as arginine, aspartic acid, glutamic acid, lysine, methionine

and mineral elements such as calcium, iron, potassium, phosphorus and zinc (Zhang et

al. 2007). In banana, several mutants have been isolated for different traits, namely

reduced height, tolerance to Fusarium wilt, early flowering, large fruit size, Black

sigatoka tolerant types (Jain, 2010b). In date palm, several mutant lines tolerant to

Bayoud disease, caused by Fusarium oxysporum f. albedinis fungus, and the plants are

already in the field for the last four years and growing well in Algeria (Jain 2007). New

mutant varieties of barley, wheat, rice and soybean with low phytic acid (LPA) have been

released and has facilitated to reduce both phosphorous pollution and increase

bioavailability of phosphorous and micronutrient minerals in cereals and legumes.

Relevance of Biotechnology and biofortification

Prasad (2010) identified three major micronutrient deficiencies in humans which include

vitamin A deficiency, iron deficiency and iodine deficiency. Micronutrient deficiency of

Zn has also received global attention (Hussain et al, 2010). In addition, improving the

content of essential amino acids in important staple foods, such as rice, has gained interest

(Welch and Graham 2004). Rice is also one of the priority crops for enhancement of the

nutritional factors such as vitamin A, Zn and iron through international schemes such as

Harvest Plus (Pfeiffer and McClafferty 2007). Increase in bioavailability of Fe in rice has

been investigated by transferring a gene for heat resistant phytase from fungal sources that

degrades phytate in plant (Bhat and Vasanthi 2005) which may also increase the Zn

bioavailability in rice.

An adequate concentration of micronutrients seems to be essentially required in major

staple crops if these crops are addressed to provide a sustainable solution to the problem

of malnutrition (Pinstrup-Anderson and Pandya- Lorch 2001). This holds true for cereals

since majority of the population in the developing world depend on cereal based food

intake. Rice alone contributes to 23% of the energy consumed worldwide and countries

that rely on rice as the main staple often consume up to 60% of their daily energy from

this cereal (Khush 2003). Conventionally, nutrient content of crops can be improved by

using field fortification strategies, to enhance the micronutrient and trace element content

of crops by applying enriched fertilizers to the soil. Biotechnological tools have generated

new opportunities to improve the amount and availability of nutrients in plant crops.

These include simple plant selection for varieties with high nutrient concentration in the

seeds, cross-breeding for incorporating a desired trait within a plant, and genetic

engineering to manipulate the nutrient content of the plant (King 2002). One of the

successful examples in using the genetic engineering approach is the production of

“Golden Rice” involving the transfer of the genes necessary for the accumulation of

Carotenoids (vitamin A precursors) in the endosperm that are not available in the rice

gene pool. The first generation Golden Rice with a gene from daffodil and a common soil

bacterium drew considerable criticism as a technological solution to a problem associated

with poverty and hunger. It was argued that Golden Rice would encourage people to rely

on a single food rather than the promotion of dietary diversification. The development of

Golden Rice 2 by replacing the daffodil gene with an equivalent gene from maize

increased the amount of beta carotene by about 20-fold resulting in about 140 grams of

the rice providing a child’s RDA for beta carotene (Raney and Pingali 2007). It has also

been recently demonstrated that beta carotene from golden rice is efficiently converted to

vitamin A in humans (Tang et al. 2009).

Neglected/underutilized crop resources for nutrition provision

Agricultural biodiversity is essential both in terms of food and nutritional security.

Diversity of kingdoms, species and gene pools can increase the productivity of farming

systems in a range of growing conditions, and more diverse farming systems are also

generally more resilient in the face of perturbations, thus enhancing food security, better

nutrition and greater health (Ochatt and Jain, 2009; Frison et al. 2011). Global food

security depends mostly on a handful of cultivated species and more than 50% of the daily

requirement of proteins and calories is derived from three major crops viz., wheat, maize

and rice (Bharucha and Pretty 2010). More than 7000 wild plant species are known to

have been used for human food at some stage in human history (Grivetti and Ogle 2000);

in India, 600 plant species are known to have food value (Rathore 2009).

In contrast, the availability of orphan- or understudied-crops as the major staple food

crops in many developing countries has contributed significantly. Some examples include,

cereals (e.g. millet, tef, fonio), legumes (cowpea, bambara groundnut, grass pea), and root

crops (cassava, yam, enset). Orphan crops are in general more adapted to the extreme soil

and climatic conditions prevalent in Africa than the major crops of the world. Minor

millets, which are high in nutrients such as calcium and iron, are grown primarily in hilly,

arid areas of India where, because of their high tolerance to drought, they are often more

productive than other grains (Tadele, 2009ab; Anon. 2010). However, due to lack of

genetic improvement, orphan crops produce inferior yield in terms of both quality and

quantity. The majority of the world’s food is produced from only a few crops, and yet

many neglected and under-utilized crops are extremely important for food production in

low income food deficit countries (LIFDCs). As the human population grows at an

alarming rate in LIFDCs, food availability has declined and is also affected due to

environmental factors, lack of improvement of local crop species, erosion of genetic

diversity and dependence on a few crop species for food supply. Neglected crops are

traditionally grown by farmers in their centres of origin or centres of diversity, where they

are still important for the subsistence of local communities, and maintained by socio-

cultural preferences and traditional uses. These crops remain inadequately characterised

and, until very recently, have been largely ignored by research and conservation.

Radiation-induced mutation techniques have successfully been used for the genetic

improvement of “major crops” and the know-how will greatly benefit genetic enhancing

of under-utilized and neglected species towards their domestication and crop

improvement. Realizing such a need, the FAO/IAEA initiated a program on genetic

improvement of under-utilized and neglected species through a Coordinated Research

Project on “Genetic Improvement of Under-utilized and Neglected Crops in LIFDCs

through Irradiation and Related Techniques” in 1998. The overall objective was to

improve food security, enhance nutritional balance, and promote sustainable agriculture in

LIFDCs (IAEA-TECHDOC.1426, 2004, Jain, 2009). The species that were studied

included medicinal and aromatic plants that are important for the West Asia and North

Africa [e.g. argel (Solenostemma arghel), caper (Capparis spp.), oregano (Origanum

syriacum), mint (Mentha piperita), liquorice (Glycyrrhiza glabra), aloe (Aloe spp.),

coriander (Coriandrum sativum), cumin (Cuminum cyminum) and henna (Lawsonia

inermis)], Andean grains for Latin America [e.g. quinoa (Chenopodium quinoa), canihua

(C. pallidicaule) and amaranth (Amaranthus caudatus)] and nutritious millets for Asia

[e.g. finger millet (Eleusine coracana), Italian millet (Setaria italica) and little millet

(Panicum miliare)] (Jain, 2009).

Reverse and forward genetics

The new gene discovery with reverse and forward genetics will open the way for

developing functional genomics plant breeding. The general strategy for reverse genetics

is called TILLING (Targeting Induced Local Lesions in Genomics) or coming together

with traditional mutagenesis functional genomics (Gilchrist and Haughn, 2005; Tadele et

al 2010). TILLING allows the identification of single-base-pair allelic variation in a target

gene in a high-throughput manner. Furthermore DNA sequence information of mutants or

crop plants facilitate the isolation of cisgenes, which are genes from crop plants

themselves or from crossable species (Jacobsen and Schouten 2010). The increasing

numbers of these isolated genes provide an opportunity to improve plant breeding while

remaining within the gene pool of the classical breeder or mutation breeder.

Finally, a multiple disciplinary approach would be ideal by including conventional and

mutation breeding together with the molecular tools for developing new crop varieties

with high yield with improved nutritional qualities in sustaining food and nutritional

security worldwide.

Conclusion

Nutrition security is integral to food security. Induced mutations are significant as novel

mutations are being isolated for enhanced nutrition quality of crop plants, for ex.

micronutrients, protein, amino acids, fatty acids and vitamins. Another source of nutrition

provision is from the neglected and underutilized crops, and requires more attention

together with the major crops for enhancing nutrition provision to the ever-growing

human population. Perhaps change of food habits would be required gradually move away

from the consumption of major crops and start using underutilized crops either singly or

in combination of both. Developing genetically novel germplasm with increased content

of these together with other health benefit components becomes more feasible concurrent

with the enhancement of breeding techniques, genomics, molecular manipulations and

genetic engineering. The cost effectiveness of applying new technologies and trained

manpower would be of paramount importance for nutrition provision to the low cost

nations.

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Table 1. Some of the major orphan crops of Africa having important nutritional

characteristics. (modified after Tadele, 2009)

Common

Name

Botanical name Important trait

African

eggplant

Solanum aethiopicum High yielding

African yam

bean

Sphenostylis stenocarpa High protein content

Amaranth Amaranthus spp. Fast growing

Bambara

groundnut

Vigna subterranea Rich in protein,

drought tolerant

Barbados

cherry

Malpighia glabra Rich in vitamin

Cassava Manihot esculentum Drought tolerant

Chickpea Cicer arietinum Protein source

Dika Irvingia gabonensis, I .

wombolu

oil-rich

Finger millet Eleusine coracana Rich in iron, protein;

low in glycaemic

index

Fonio Digitaria exilis Fast maturing

Noug Guizotia abyssinica High oil content

Quinoa Chenopodium quinoa High in

protein content

Sesame Sesamum indicum oxidatively stable oil

Sweet potato Ipomoea batatas rich in riboflavin and

calcium

Tef Eragrostis tef Tolerant to abiotic

stresses; free of gluten

Vernonia Vernonia galamensis High oil content

Figure 1. Global prevalence of micronutrient malnutrition* (Ramakrishnan 2002)

*For more details, please refer Ramakrishnan (2002)

Figure 2: Differential response of citrus varieties to different doses of gamma radiation

treatment. Citrus var. Losslille is more radiation tolerant when compared to two other

varieties: 30 Gy dose promotes shoot growth, which is much better than the control as well as

other two varieties.

Radiation dose effect on

citrus

Figure 3. Global mutant varieties with nutritional quality and other desirable attributes (based on the data from www-mvd.iaea.org; September 2010)

Figure 4: A new sweet sorghum mutant variety, which is suitable for food, feed, and bio-

energy

New Sweet Sorghum Mutant Variety Yuantian No.1