Breeding for micronutrients in staple food crops from a human nutrition perspective

Over three billion people are currently micronutrient (i.e. micronutrient elements and vitamins) malnourished, resulting in egregious societal costs including learning disabilities among children, increased morbidity and mortality rates, lower worker productivity, and high healthcare costs, all factors diminishing human potential, felicity, and national economic development. Nutritional deficiencies (e.g. iron, zinc, vitamin A) account for almost two-thirds of the childhood death worldwide. Most of those afflicted are dependent on staple crops for their sustenance. Importantly, these crops can be enriched (i.e. 'biofortified') with micronutrients using plant breeding and/or transgenic strategies, because micronutrient enrichment traits exist within their genomes that can to used for substantially increasing micronutrient levels in these foods without negatively impacting crop productivity. Furthermore, 'proof of concept' studies have been published using transgenic approaches to biofortify staple crops (e.g. high beta-carotene 'golden rice' grain, high ferritin-Fe rice grain, etc). In addition, micronutrient element enrichment of seeds can increase crop yields when sowed to micronutrient-poor soils, assuring their adoption by farmers. Bioavailability issues must be addressed when employing plant breeding and/or transgenic approaches to reduce micronutrient malnutrition. Enhancing substances (e.g. ascorbic acid, S-containing amino acids, etc) that promote micronutrient bioavailability or decreasing antinutrient substances (e.g. phytate, polyphenolics, etc) that inhibit micronutrient bioavailability, are both options that could be pursued, but the latter approach should be used with caution. The world's agricultural community should adopt plant breeding and other genetic technologies to improve human health, and the world's nutrition and health communities should support these efforts. Sustainable solutions to this enormous global problem of 'hidden hunger' will not come without employing agricultural approaches.     

Exploiting genotypic variation in plant nutrient accumulation to alleviate micronutrient deficiency in populations

More than 2 billion people consume diets that are less diverse than 30 years ago, leading to deficiencies in micronutrients, especially iron (Fe), zinc (Zn), selenium (Se), iodine (I), and also vitamin A. A strategy that exploits genetic variability to breed staple crops with enhanced ability to fortify themselves with micronutrients (genetic biofortification) offers a sustainable, cost-effective alternative to conventional supplementation and fortification programs. This is more likely to reach those most in need, has the added advantages of requiring no change in current consumer behaviour to be effective, and is transportable to a range of countries. Research by our group, along with studies elsewhere, has demonstrated conclusively that substantial genotypic variation exists in nutrient (e.g. Fe, Zn) and nutrient promotor (e.g. inulin) concentrations in wheat and other staple foods. A rapid screening technique has been developed for lutein content of wheat and triticale, and also for pro-vitamin A carotenoids in bread wheat. This will allow cost-effective screening of a wider range of genotypes that may reveal greater genotypic variation in these traits. Moreover, deeper understanding of genetic control mechanisms and development of molecular markers will facilitate breeding programs. We suggest that a combined strategy utilising plant breeding for higher micronutrient density; maximising the effects of nutritional promoters (e.g. inulin, vitamin C) by promoting favourable dietary combinations, as well as by plant breeding; and agronomic biofortification (e.g. adding iodide or iodate as fertiliser; applying selenate to cereal crops by spraying or adding to fertiliser) is likely to be the most effective way to improve the nutrition of populations. Furthermore, the importance of detecting and exploiting beneficial interactions is illustrated by our discovery that in Fe-deficient chickens, circulating Fe concentrations can be restored to normal levels by lutein supplementation. Further bioavailability/bioefficacy trials with animals and humans are needed, using varying dietary concentrations of Fe, Zn, carotenoids, inulin, Se and I to elucidate other important interactions in order to optimise delivery in biofortification programs.     

Potential impact and cost-effectiveness of multi-biofortified rice in China

Biofortification, that is, improving the micronutrient content of staple foods through crop breeding, could be a pro-poor, pro-rural, agriculture-based intervention to reduce the health burden of micronutrient malnutrition. While the potential cost-effectiveness of crops biofortified with single micronutrients was shown in previous research, poor people often suffer from multiple micronutrient deficiencies, which should be accounted for in biofortification initiatives. This study is the first to estimate the potential health benefits and cost-effectiveness of multi-biofortification. Rice with enhanced provitamin A, zinc, iron and folate concentrations is used as a concrete example. The research is conducted for China, the largest rice producer in the world, where micronutrient malnutrition remains a major public health problem. Using the DALY (disability-adjusted life year) framework, the current annual health burden of the four micronutrient deficiencies in China is estimated at 10.6 million DALYs. Introducing multi-biofortified rice could lower this burden by up to 46%. Given the large positive health impact and low recurrent costs of multi-biofortification, this intervention could be very cost effective: under optimistic assumptions, the cost per DALY saved would be around US$ 2; it would stay below US$ 10 even under pessimistic assumptions.     

Biofortified crops to alleviate micronutrient malnutrition

Micronutrient malnutrition affects more than half of the world population, particularly in developing countries. Concerted international and national fortification and supplementation efforts to curb the scourge of micronutrient malnutrition are showing a positive impact, alas without reaching the goals set by international organizations. Biofortification, the delivery of micronutrients via micronutrient-dense crops, offers a cost-effective and sustainable approach, complementing these efforts by reaching rural populations. Bioavailable micronutrients in the edible parts of staple crops at concentrations high enough to impact on human health can be obtained through breeding, provided that sufficient genetic variation for a given trait exists, or through transgenic approaches. Research and breeding programs are underway to enrich the major food staples in developing countries with the most important micronutrients: iron, provitamin A, zinc and folate.     

Biotechnology for Enhanced Nutritional Quality in Plants

With almost 870 million people estimated to suffer from chronic hunger worldwide, undernourishment represents a major problem that severely affects people in developing countries. In addition to undernourishment, micronutrient deficiency alone can be a cause of serious illness and death. Large portions of the world population rely on a single, starch-rich crop as their primary energy source and these staple crops are generally not rich sources of micronutrients. As a result, physical and mental health problems related to micronutrient deficiencies are estimated to affect around two billion people worldwide. The situation is expected to get worse in parallel with the expanding world population. Improving the nutritional quality of staple crops seems to be an effective and straightforward solution to the problem. Conventional breeding has long been employed for this purpose but success has been limited to the existing diversity in the gene pool. However, biotechnology enables addition or improvement of any nutrient, even those that are scarce or totally absent in a crop species. In addition, biotechnology introduces speed to the biofortification process compared to conventional breeding. Genetic engineering was successfully employed to improve a wide variety of nutritional traits over the last decade. In the present review, progress toward engineering various types of major and minor constituents for the improvement of plant nutritional quality is discussed.     

Effect of different cooking methods on loss of iron and zinc micronutrients in fortified and non-fortified rice

Malnutrition is considered as major public health concern and is emerging challenge to food and nutrition security particularly in developing countries. Rice is the staple food and consumed by the half of the world's population which is the source of daily requirement of the nutrients. Attempts are being made to fortify rice with micronutrients, but the loss or retention of these micronutrients in different cooking methods is not well studied and documented especially in fortified rice. In the present study, paddy seeds of six Indian varieties were fortified with iron and zinc by parboiling process. Consequently, fortified polished rice had higher micronutrient contents (Fe, 106.31 ± 12.56; Zn, 97.72 ± 9.75) than non-fortified polished rice (Fe, 7.44 ± 1.05; Zn, 14.74 ± 2.94) expressed in ppm. Polished rice of both fortified and non-fortified were cooked under five different cooking conditions and analyzed for remaining iron and zinc content. Cooking rice in rice cooker without prior washing (NRC) retained highest concentration of Fe and Zinc in both fortified and non-fortified rice varieties. It also showed that fortified rice suffered higher percentage loss of micronutrient, than the non-fortified rice. But the average retained micronutrient amount measured in ppm, was higher in fortified rice (Fe, 43.54 ± 6.88; Zn, 36.7 ± 3.12) than in non-fortified rice (Fe, 4.24 ± 0.87; Zn, 9.3 ± 2.11). Hence, adopting appropriate cooking method, higher amount of micronutrients will be retained in the cooked food which will in turn help in combating the malnutrition and improve health.     

小麦子粒铁、锌元素含量的研究进展

目前,铁、锌元素营养失衡已严重影响人类健康,而小麦作为人们的主要食物,其子粒中铁、锌元素含量普遍较低。因此,通过小麦育种策略提高铁、锌元素含量则被认为是最经济有效的手段,这也引起了国内外学者的广泛关注。本文概述了小麦子粒铁、锌元素含量的遗传差异、影响因素以及分子机理的研究现状,并展望了提高小麦子粒铁、锌元素含量的研究前景及途径。     

The impact of mineral nutrients in food crops on global human health

Nutrient sufficiency is the basis of good health, productive lives and longevity for everyone. Nutrient availability to people is primarily determined by the output of foods produced from agricultural systems. If agricultural systems fail to provide enough food diversity and quantity to satisfy all the nutrients essential to human life, people will suffer, societies will deteriorate and national development efforts will stagnate. Importantly, plant foods provide most of the nutrients that feed the developing world. Unfortunately, as a result of population pressures, many global food systems are not currently providing enough micronutrients to assure adequate micronutrient intakes for all people. This has resulted in an increasing prevalence of micronutrient deficiencies (e.g., iron deficiency, vitamin A deficiency, and iodine deficiency disorders) that now afflicts over three billion people globally mostly among resource-poor women, infants and children in developing countries. The consequences of micronutrient malnutrition are profound and alarming for human existence. Agricultural approaches to finding sustainable solutions to this problem are urgently needed. This review presents some ways in which plant nutritionists can contribute to preventing micronutrient malnutrition in sustainable ways.     

Micronutrient status and fertilizer use scenario in India

India was dependent on external food supplies in the early 1960s. To meet the growing demand for food, fiber and fuel, high yielding cultivars were introduced. These high yielding crop cultivars were highly responsive to fertilizers. Thus, slowly the soils were exhausted of their nutrients. Application of major nutrients (nitrogen, phosphorus, potassium) became common, therefore the crops started responding to micronutrient fertilizers. Concerted efforts have been made through the All India Coordinated Research Project on Micronutrients to delineate the soils of India regarding the deficiency of micronutrients. At present about 48.1% of Indian soils are deficient in diethylenetriaminepentaacetate (DTPA) extractable zinc, 11.2% in iron, 7% in copper and 5.1% in manganese. Apart from the deficiency of these micronutrients, deficiencies of boron and molybdenum have also been reported in some areas. Areas with multi-micronutrient deficiencies are limited, thus simple fertilizers are sufficient to exploit the potential of crops and cropping systems. Based on the extent of deficiency, cultivated area, and crop removal, the micronutrient fertilizer demand in 2025 is projected using sufficiency and maintenance approaches.     

Micronutrient status and fertilizer use scenario in India

India was dependent on external food supplies in the early 1960s. To meet the growing demand for food, fiber and fuel, high yielding cultivars were introduced. These high yielding crop cultivars were highly responsive to fertilizers. Thus, slowly the soils were exhausted of their nutrients. Application of major nutrients (nitrogen, phosphorus, potassium) became common, therefore the crops started responding to micronutrient fertilizers. Concerted efforts have been made through the All India Coordinated Research Project on Micronutrients to delineate the soils of India regarding the deficiency of micronutrients. At present about 48.1% of Indian soils are deficient in diethylenetriaminepentaacetate (DTPA) extractable zinc, 11.2% in iron, 7% in copper and 5.1% in manganese. Apart from the deficiency of these micronutrients, deficiencies of boron and molybdenum have also been reported in some areas. Areas with multi-micronutrient deficiencies are limited, thus simple fertilizers are sufficient to exploit the potential of crops and cropping systems. Based on the extent of deficiency, cultivated area, and crop removal, the micronutrient fertilizer demand in 2025 is projected using sufficiency and maintenance approaches.     

Biofortification of crops for reducing malnutrition

Micronutrient deficiencies affect approximately 3 billion people worldwide. Malnutrition hinders the development of human potential and social and economic development in developing countries. The World Health Organization (WHO) and the Consultative Group on International Agricultural Research (CGIAR) have made fighting micronutrient deficiencies, known as hidden hunger, a high priority. Deficiencies of the micronutrients, such as iron, zinc, and vitamin A, are the most devastating among the world’s poor. WHO emphasizes nutrient supplementation and food fortification to address the malnutrition. CGIAR has placed a greater emphasis on biofortification through the HarvestPlus challenge program, and improved micronutrient content of the staple crops (rice, wheat, maize, beans, cassava, pearl millet, and sweet potato) through breeding and biotechnological approaches. An excellent example of biotechnology application is the development of ‘golden rice’ with adequate levels of a provitamin A, β-carotene. The Africa Harvest and the BioCassava Plus programs, respectively, are developing sorghum and cassava with improved nutritional quality. Here, we summarize current strategies of crop biofortification and future prospects towards the development of biofortified crops.     

Seed priming with iron and zinc in bread wheat: effects in germination,mitosis and grain yield

Currently, the biofortification of crops like wheat with micronutrients such as iron (Fe) and zinc (Zn) is extremely important due to the deficiencies of these micronutrients in the human diet and in soils. Agronomic biofortification with Fe and Zn can be done through different exogenous strategies such as soil application, foliar spraying, and seed priming. However, the excess of these micronutrients can be detrimental to the plants. Therefore, in the last decade, a high number of studies focused on the evaluation of their phytotoxic effects to define the best strategies for biofortification of bread wheat. In this study, we investigated the effects of seed priming with different dosages (1 mg L^?1 to 8 mg L^?1) of Fe and/or Zn in germination, mitosis and yield of bread wheat cv. ‘Jordão’ when compared with control. Overall, our results showed that: micronutrient dosages higher than 4 mg L^?1 negatively affect the germination; Fe and/or Zn concentrations higher than 2 mg L^?1 significantly decrease the mitotic index and increase the percentage of dividing cells with anomalies; treatments performed with 8 mg L^?1 of Fe and/or 8 mg L^?1 Zn caused negative effects in germination, mitosis and grain yield. Moreover, seed priming with 2 mg L^?1 Fe?+?2 mg L^?1 Zn has been shown to be non-cytotoxic, ensuring a high rate of germination (80%) and normal dividing cells (90%) as well as improving tillering and grain yield. This work revealed that seed priming with Fe and Zn micronutrients constitutes a useful and alternative approach for the agronomic biofortification of bread wheat.     

Golden Rice and ‘Golden’ crops for human nutrition

Micronutrients are essential for a healthy life. Humans do not produce micronutrients, and hence they must obtain them through the foodchain. Staple crops are the predominant food source of mankind, but need to be complemented by other foodstuffs because they are generally deficient in one or the other micronutrient. Breeding for micronutrient-dense crops is not always a viable option because of the absence of genetic variability for the desired trait. Moreover, sterility issues and the complex genetic makeup of some crop plants make them unamenable to conventional breeding. In these cases, genetic modification remains the only viable option. The tools to produce a number of micronutrients in staple crops have recently become available thanks to the identification of the genes involved in the corresponding biochemical pathways at an unprecedented rate. Discarding genetic modification as a viable option is definitely not in the interest of human wellbeing.     

Global impacts of human mineral malnutrition

Malnutrition—in the form of insufficient energy intakes—affects millions of people worldwide and the negative impact of this kind of hunger is well acknowledged, not least by agronomists trying to increase yields to ensure a sufficient supply of food. This review focuses on another, more particular and “hidden” form of malnutrition, namely mineral malnutrition. It illustrates the burden of disease that is caused by mineral deficiencies and the social and economic consequences they bring about. Mineral malnutrition has a considerable negative impact on individual well-being, social welfare and economic productivity. Agricultural scientists should keep the nutritional qualities of food in mind and—next to optimizing the agricultural properties of crops that are paramount for their adoption by farmers—in particular try to increase the micronutrient content in major staple crops as one way to address vitamin and mineral malnutrition in humans; especially plant breeding approaches promise to be very cost-effective.     

Zinc,Iron, Manganese and Copper Uptake Requirement in Response to Nitrogen Supply and the Increased Grain Yield of Summer Maize

The relationships between grain yields and whole-plant accumulation of micronutrients such as zinc (Zn), iron (Fe), manganese (Mn) and copper (Cu) in maize (Zea mays L.) were investigated by studying their reciprocal internal efficiencies (RIEs, g of micronutrient requirement in plant dry matter per Mg of grain). Field experiments were conducted from 2008 to 2011 in North China to evaluate RIEs and shoot micronutrient accumulation dynamics during different growth stages under different yield and nitrogen (N) levels. Fe, Mn and Cu RIEs (average 64.4, 18.1and 5.3 g, respectively) were less affected by the yield and N levels. ZnRIE increased by 15% with an increased N supply but decreased from 36.3 to 18.0 g with increasing yield. The effect of cultivars on ZnRIE was similar to that of yield ranges. The substantial decrease in ZnRIE may be attributed to an increased Zn harvest index (from 41% to 60%) and decreased Zn concentrations in straw (a 56% decrease) and grain (decreased from 16.9 to 12.2 mg kg⁻¹) rather than greater shoot Zn accumulation. Shoot Fe, Mn and Cu accumulation at maturity tended to increase but the proportions of pre-silking shoot Fe, Cu and Zn accumulation consistently decreased (from 95% to 59%, 90% to 71% and 91% to 66%, respectively). The decrease indicated the high reproductive-stage demands for Fe, Zn and Cu with the increasing yields. Optimized N supply achieved the highest yield and tended to increase grain concentrations of micronutrients compared to no or lower N supply. Excessive N supply did not result in any increases in yield or micronutrient nutrition for shoot or grain. These results indicate that optimized N management may be an economical method of improving micronutrient concentrations in maize grain with higher grain yield.     

Plant nutrition research: Priorities to meet human needs for food in sustainable ways

The world population is expanding rapidly and will likely be 10 billion by the year 2050. Limited availability of additional arable land and water resources, and the declining trend in crop yields globally make food security a major challenge in the 21st century. According to the projections, food production on presently used land must be doubled in the next two decades to meet food demand of the growing world population. To achieve the required massive increase in food production, large enhancements in application of fertilizers and improvements of soil fertility are indispensable approaches. Presently, in many developing countries, poor soil fertility, low levels of available mineral nutrients in soil, improper nutrient management, along with the lack of plant genotypes having high tolerance to nutrient deficiencies or toxicities are major constraints contributing to food insecurity, malnutrition (i.e., micronutrient deficiencies) and ecosystem degradation. Plant nutrition research provides invaluable information highly useful in elimination of these constraints, and thus, sustaining food security and well-being of humans without harming the environment. The fact that at least 60% of cultivated soils have growth-limiting problems with mineral-nutrient deficiencies and toxicities, and about 50% of the world population suffers from micronutrient deficiencies make plant nutrition research a major promising area in meeting the global demand for sufficient food production with enhanced nutritional value in this millennium. Integration of plant nutrition research with plant genetics and molecular biology is indispensable in developing plant genotypes with high genetic ability to adapt to nutrient deficient and toxic soil conditions and to allocate more micronutrients into edible plant products such as cereal grains.     

Critical evaluation of strategies for mineral fortification of staple food crops

Staple food crops, in particular cereal grains, are poor sources of key mineral nutrients. As a result, the world’s poorest people, generally those subsisting on a monotonous cereal diet, are also those most vulnerable to mineral deficiency diseases. Various strategies have been proposed to deal with micronutrient deficiencies including the provision of mineral supplements, the fortification of processed food, the biofortification of crop plants at source with mineral-rich fertilizers and the implementation of breeding programs and genetic engineering approaches to generate mineral-rich varieties of staple crops. This review provides a critical comparison of the strategies that have been developed to address deficiencies in five key mineral nutrients—iodine, iron, zinc, calcium and selenium—and discusses the most recent advances in genetic engineering to increase mineral levels and bioavailability in our most important staple food crops.     

The contribution of transgenic plants to better health through improved nutrition: opportunities and constraints

Malnutrition is a prevalent and entrenched global socioeconomic challenge that reflects the combined impact of poverty, poor access to food, inefficient food distribution infrastructure, and an over-reliance on subsistence mono-agriculture. The dependence on staple cereals lacking many essential nutrients means that malnutrition is endemic in developing countries. Most individuals lack diverse diets and are therefore exposed to nutrient deficiencies. Plant biotechnology could play a major role in combating malnutrition through the engineering of nutritionally enhanced crops. In this article, we discuss different approaches that can enhance the nutritional content of staple crops by genetic engineering (GE) as well as the functionality and safety assessments required before nutritionally enhanced GE crops can be deployed in the field. We also consider major constraints that hinder the adoption of GE technology at different levels and suggest policies that could be adopted to accelerate the deployment of nutritionally enhanced GE crops within a multicomponent strategy to combat malnutrition.     

Nitrate assimilation in rice seedlings as affected by mineral nutrition

Deficiencies of each macronutrient (N, P, K, Ca, Mg and Fe)in the culture solution depressed the specific activities ofnitrate reductase (NR) and nitrite reductase (NiR) from riceseedlings. Nitrate and potassium deficiencies especially loweredNR induction, whereas phosphorus deficiency caused the leastdecrease in enzyme induction. On the other hand the activityof NiR was decreased most by deficiencies of nitrate and phosphorus.Potassium deficiency was not as effective in suppressing theinduction of NiR. Sulfur deficiency slightly promoted the inductionof both NR and NiR. Generally, micronutrient deficiencies didnot affect either enzyme. NR induction was slightly decreasedby B, Zn, Cu and Mo deficiencies, and increased by Mn deficiency;whereas NiR activity was slightly increased by B and Cu deficiencies,and was not affected by other micronutrients. Nitrate contentwas decreased by deficiencies of N, P, K, Ca, and micronutrients,and unaffected by Mg, Fe and S deficiencies. Glutamic acid dehydrogenase(GDH) activity was increased by N, Fe and P deficiencies, anddecreased by Mo and Zn deficiencies, and unaffected by othernutrient treatments. (Received August 25, 1976; )     

Nutritional enhancement of rice for human health: The contribution of biotechnology

Micronutrient malnutrition is widespread, especially in poor populations across the globe where daily caloric intake is confined mainly to staple cereals. Rice, which is a staple food for over half of the world's population, is low in bioavailable micronutrients required for the daily diet. Improvements of the plant-based diets are therefore critical and of high economic value in order to achieve a healthy nutrition of a large segment of the human population. Rice grain biofortification has emerged as a strategic priority for alleviation of micronutrient malnutrition. Nutritional enhancement of crops through conventional breeding is often limited by the low genetic variability for required dietary micronutrient levels. In this case, biotechnology strategies offer effective and efficient perspectives. In this review, we discuss genetic engineering approaches that have been successful in the nutritional enhancement of rice endosperm. These advancements will make substantial contributions to crop improvement and human nutrition. Their practical application, however, also demands visionary changes in regulatory policies and a broader consumer acceptance.