eChapters

The hybrid breeding program in pigeonpea is unique and the first among the food legume. The concept of hybrid in this crop was conceived in 1974; and it is based on a small window of natural out-crossing present in the crop. This natural out-crossing in pigeonpea is mediated by a range of flying insects, which forage on large bright colored flowers of the crop in search of nectar and in the process affect the cross-fertilization. In the last 40 years a lot of basic and applied research has been successfully completed and over 50 research papers have been published in national and international journals of repute. A coordinated effort of scientists of different institutions has led to the release of two hybrids, one for central and another for peninsular zone of India.

With the advent of hybrid technology, it is natural to assess its potential benefits at the national level. To meet the domestic requirement of pigeonpea, about 50.000–60,000 metric tons of pigeonpea is imported annually from Myanmar and Africa. In this communication, an attempt has been made to look into the possibility if we can meet the entire domestic needs of pigeonpea through the adoption of hybrid technology. It is a big question but the goal is not impossible. The estimates shown in Table 18 show that if only 2% of total pigeonpea area can be brought under hybrids, the target can be met with ease. This will require seed production in only 700-800 hectares.

To overcome the challenge of popularizing pigeonpea hybrids in the country, it is considered fair enough to compare the hybrid rice with respect to their seed production technology, yield advantages and profitability to the farmers and seed producers. Since both pigeonpea and rice are grouped under self-pollinated crops, this comparison, in the views of authors, may help in encouraging hybrid pigeonpea seed producers and those involved in marketing of hybrid seed.
  
Rice is a highly self-pollinated crop and earlier hybrids were not possible because it was essential that large quantities of hybrid seeds are produced easily and economically. In China, the birth place of hybrid rice, this impossible looking task was accomplished. To launch this program, the Chinese scientists decided to alter the pollination system of rice.

Fig. 25: ICPH 2671 in field

5.1 Seed back up programs

In hybrid breeding program, testing of new hybrid combinations is a continuous process and in most cases, the availability of crossed (hybrid) seed for their annual evaluation of selected hybrids is an important issue. Since, the selection of hybrids for advancement is based on yield data, one year will be required to produce seed for testing in the subsequent season. This means for each cycle of advancement, one valuable season will be consumed in their seed production. To avoid this situation and enhance the testing program at a fast pace, an aggressive hybrid seed production program should be followed. This can be achieved by visually selecting relatively more number of hybrids based on their flowering time, plant vigor, disease resistance, and early pod load about 4-5 weeks before harvesting. During this period, the new seed of the selected hybrids can be produced manually. This operation is facilitated by the perennial nature of pigeonpea plant, which can flower for extended periods of 40-50 days; that allows completion of desired number of pollinations of the target crosses in the same cropping season. One trained person can pollinate about 300 buds in 6-7 hours; and with 20% success, about 150 seeds can be produced. Experience has shown that, with good planning and crop management, this program can be achieved with ease.

4.1. Flower and flowering in pigeonpea

Growth habit: Primarily, there are two major growth types in pigeonpea. These are determinate and non-determinate. The determinate growth in the pigeonpea plants is characterized by flowers in bunches at the top of the canopy (Fig 17a). In this case the epical growing points are transformed in to reproductive buds and bunches of flowers appear on the top of the canopy. Such cultivars are not popular as the management of insects such as Helicoverpa and Maruca becomes very difficult. Most of the cultivated types belong to another growth category called as non-determinate. In such genotypes the apical growing points remain vegetative and the plants keep growing under favorable conditions and it results in tall canopy with many spreading or compact branches. The inflorescence in this case emerges from the axil of secondary and tertiary branches (Fig 17b); and hang sideways. These types show less susceptibility to insects due to non-clustering nature of flowers.

3.1. Natural cross-pollination

Breeding procedures in any plant species of economic importance are guided by its reproductive system particularly, pollination mechanism. In the cereal crops such as maize, pearl millet etc., where cross-pollination is facilitated by wind, the breeders have developed schemes for commercial exploitation of dominance and epistatic genetic variation through hybrid and population breeding. In contrast in self-pollinated crops, the efforts are always made to exploit additive genetic variation primarily through pedigree and mass selection.
 
In pulses, self-pollination is a rule and pigeonpea is no exception. However under field conditions, occurrence of hybrid plants in the populations is not uncommon; and it happens due to some degree of natural out-crossing that took place in the preceding generation. This type of partial natural out-crossing in pigeonpea is affected by a variety of insects (Fig 10) which forage on its large flowers to collect the nectar. These minor incidences of out-crossing are the primary source of varietal deterioration. Recently, the pigeonpea breeders converted this constraint into an opportunity for genetic enhancement of yield through exploitation of hybrid vigor and developing commercial hybrids.

Male sterility is a situation where the male reproductive parts of a plant are either absent, aborted, or non-functional; and therefore, they fail to participate in the process of natural sexual reproduction. This situation generally arises due to some developmental defect that occurs at any stage of microsporogenesis (process of development or release of pollen grains) in the plants. Since male sterility is the manifestation of abnormal growth and development and can occur at any point of time, the action of genes controlling male sterility is also be variable and inconsistent across the genotypes. Based on the type of malfunctioning of androecium, the male sterility systems have been classified as structural (absence or deformity of anthers), sporogenous (defective microsporogenesis), and functional (failure of mature pollen to germinate). In addition, on the basis of genetic control mechanism of the male sterility, it has also been classified as genetic, cytoplasmic, and cytoplasmic nuclear (or genetic) male sterility. With respect to utilization of male sterility in crop breeding, it is essential that individuals with altered male fertility keep their female fertility intact. The fertilization of such plants takes place with pollen grains from other plant that is transferred with the help of external agencies such as wind, insects, or human beings.

Pigeonpea (Cajanus cajan L. Millsp.) is crowned as poor man’s crop which provides much needed protein to farming families with least inputs. It is a versatile plant species which can grow successfully in a range of soil types, temperatures and photoperiods. Its deep root system enables it to overcome intermittent drought and other stresses. In India, the crop is grown annually on about 4.04 m ha (FAO, 2012) and the major pigeonpea growing states are Maharashtra, Andhra Pradesh, Karnataka, Madhya Pradesh, Uttar Pradesh, Bihar and Gujarat (Table 1). The high protein pigeonpea dal (decorticated dry splits) is widely consumed across the country in various cuisines. The annual national production of pigeonpea is about 2.56 m tones. This produce, however, is not sufficient to feed the ever growing population of the country, and hence, necessitates huge (500,000 tons) imports of this pulse. In spite of its importance in food security and sincere research efforts, its national productivity could never cross the bar of 800 kg/ha (Fig 1) since independence. Since last 50 years, ICAR has been very seriously pursuing the issue of genetic enhancement with huge investments and some out-standing pure line varieties with about 10% more grains over the best local varieties were developed.

Abstract

Nitrogen is the most limiting nutrient controlling production of all agricultural systems. Application of N fertilizer has increased food production tremendously all over the world. However, it also contributes towards global warming and ozone layer depletion through emission of nitrous oxide, a greenhouse gas (GHG). Therefore, there is a need to assess the possibilities of N management for climate change abatement and food security enhancement, while minimizing environmental and human health impacts. In this chapter, role of gaseous-N emission on N use and various mitigation options for Indian agriculture have been discussed. For the base year 2007, the direct and indirect N₂O emissions from Indian agricultural soils was estimated to be 136.29 Gg (35.3 Tg CO₂ equivalent) and 30.61 Gg (5.8 Tg CO₂ equivalent), respectively.

Abstract

Increases in atmospheric concentrations of CO₂ are likely to have major physiological consequences for plants, due to the major effects of atmospheric CO₂ on photosynthetic rates and the centrality of photosynthesis to plant metabolism. We review what is known about the effects of growth at elevated levels of CO₂ on the protein and N concentrations of plants in general, and food crops in particular. Growth at elevated CO₂ typically reduces plant tissue N concentrations by 10-15%. The physiological mechanisms behind this decrease in tissue protein and N are likely complex, and include both dilution from increased tissue concentrations of non-structural carbohydrates and decreased root uptake of N.

Abstract

During the past five decades, the role of nitrogen (N) fertilizer in augmenting foodgrain production has widely been recognized the world over. In Indian soils, N deficiency is recognized as a universal problem and its consumption has increased by more than 300 times during the last 60 years.

Abstract

Intensification of agriculture during the last few decades has resulted in considerable depletion of the productivity of Indian soils and the balance sheet of potassium (K) is highly negative due to very low level of K fertilization. Although the need for balanced fertilization is known for many years, it is not always observed in practice by farmers. Since the effects of nitrogen are easily seen and crops look more vigorous and yield better, excessive use of nitrogenous fertilizers, accompanied by an inadequate supply of other nutrients, is considered by farmers as a reasonable insurance against yield losses. Often, the lack of K restricts nitrogen uptake and utilization, and hence plant growth. When K is limiting, nitrogen fertilizers are used inefficiently by the plants. This nutrient imbalance is thus closely associated with low nitrogen-use-efficiency (NUE) of crop plants, which is detrimental to crop production and environment – because of the consequences of N unused by the crop.

Abstract

The abiotic stresses viz. drought, high temperature, water stress, salinity, heavy metal contaminations etc. cause variable effect on nitrogen (N) mineralization, its uptake and transport, assimilation and remobilization, and subsequently the growth and productivity of plants. The affect is generally negative, however, the magnitude of decrease in various processes is dependent on genotype, age of the plant, tissue under investigation, type and extent of the stress and other agro-climatic conditions. Biological nitrogen fixation and nitrate assimilation appear to be more sensitive than the ammonium assimilation possibly because re-assimilation of ammonia released in the tissue due to catabolism of N molecules is essential for the survival of plants. In addition, synthesis of stress metabolites (proline, glycine betaine and poly amines) is directly dependent on the availability of primary amino acid, glutamic acid. There are few well coordinated and detailed studies, determining the key factors involved in regulating NUE under abiotic stresses.

Abstract

Nitrogen is the most essential nutrient for all plants and is also the major limiting factor in plant productivity. High yielding varieties of all crops are responsive to nitrogen but its ever-increasing use has shown detrimental impact on the environment along with extremely low use efficiency. Nitrogen use efficiency (NUE) in crop plants depends on various internal and external factors, which are dealt in detail. There has been a significant interest in genetic engineering of the crops to improve NUE. The use of biotechnological interventions by manipulating genes of the nitrogen utilization pathway to improve NUE has not been very successful. But transgenics/mutants with modified capacities for nitrate uptake, assimilation and remobilization have enhanced our understanding of the genetic control of NUE. In both cellular, and at whole plant level the mechanisms involved in N remobilization from the senescing organs towards the grain have recently gained importance but their understanding is still preliminary. Recent evidences have shown that grain filling during later stages of crop growth is supported by N recycling.

Abstract

Nitrogen is the most important element in terms of nutrition for plants, nitrate being the most preferred form. Nitrate not only acts as nutritional element but also as a signal to modulate metabolism and plant architecture. Nitrate sensing mostly happens in the root tip through a hitherto unknown mechanism, but its response can be demonstrated on an organism-wide scale. Functional genomic studies revealed over a thousand nitrate-responsive genes, involved not only in N and C metabolism, but also in various other physio-logical processes. Identification of nitrate response element/s common to all these genes could pave the way to unravel the mechanism of nitrate signaling, but findings in this direction have remained inconclusive so far. Several trans-acting factors have been implicated in N signaling and response, but none of these have been convincingly demonstrated to be specific to this response. Nitrate uptake, which is sometimes associated with nitrate sensing, is also highly regulated process and involves multiple transport systems like HATS, LATS and dual affinity transporters.

Abstract

Nitrate reductase (NR) catalyzes the reduction of nitrate to nitrite, which is the first step in the nitrate assimilation pathway, but can also reduce nitrite to nitric oxide (NO), an important signaling molecule that is thought to mediate a wide array of developmental and physiological processes in plants. Accordingly, the amount of NR protein and its catalytic activity are closely controlled in the cell. This chapter focuses on the posttranslational regulation of NR activity, which has been known for some time to involve reversible protein phosphorylation. The hinge 1 region of NR contains the serine-534 regulatory site, which when phosphorylated can bind a 14-3-3 protein resulting in inhibition of NR activity. This fundamental mechanism underlies the rapid modulation of NR activity in leaves in response to light/dark transitions, and the activation of NR that occurs in response to hypoxia (e.g., as with soil flooding). Recent results suggest that the basis for hypoxia-induced NR activation, which is thought to be essential for plant survival, may involve reactive oxygen species (ROS), such as H₂O₂, that are produced in response to many plant stressors including hypoxia.

Abstract

Nitrate uptake from the soil is the first step in nitrate acquisition and utilization by plants. Nitrate uptake and its release into cells is mediated by nitrate transporter(s) located into the plasma membrane of the root. Three transporters have been identified by kinetic measurements in plant roots. These are constitutive high affinity nitrate transporter (cHATS), inducible high affinity nitrate transporter (iHATS) and low affinity nitrate transporter (LATS). Two of these display saturable kinetics; a low capacity constitutive system and a high capacity inducible system. In addition, a non-saturating low affinity, high capacity system becomes apparent only at higher external NO^-₃ concentration. Genes that encode representatives of each class of transport system have been identified and fall into two families: NRT1 and NRT2. Members of these families are induced in response to nitrate in the environment and are regulated by internal signals including nitrogen metabolites and shoot demand for nitrogen. This chapter deals with the physiology and molecular biology of nitrate transporters in higher plants.

Introduction

Engineered nanoparticles are a group of products that are defined by their size, ranging from 1 to 100 nm. Due to the smaller size, the physio-chemical properties of nano particles become very different from those of the same material in a larger form. Total production of nano-materials is expected to increase considerably in the near future. Nanotoxicology is yet another emerging field of science dealing with the effects of nano-devices and engineered nano-particles on living organisms. It has been observed that the materials in nano-dimensions react totally different in comparison to their existence in micro or macro sizes.

Introduction

The characterization of the potential health impact of engineered materials produced through nanotechnology is an emerging issue of considerable discussion and debate. The current focus on nanotoxicity fails to assess the risk in different local environments and populations. People in developing countries may be more prone to adverse effects of nanoparticles because of underlying health conditions and malnutrition. Moreover, genetic susceptibility to toxic effects varies in diverse ethnic groups and geographical areas. The scientific community needs to identify these information gaps before developing regulations and standard methodologies for nanotoxicity assessment.