Genetic Enhancement of Rabi Sorghum: Adapting the Indian Durras

Genetic Enhancement of Rabi Sorghum – Adapting the Indian Durras

P. Sanjana Reddy

Directorate of Sorghum Research, Rajendranagar, Hyderabad, India

J.V. Patil

Directorate of Sorghum Research, Rajendranagar, Hyderabad, India

Table of Contents

Cover image

Title page

Copyright

Preface

Chapter 1. Introduction

1.1 Production Statistics

1.2 Environmental Factors Limiting Rabi Sorghum Productivity

1.3 Uses

1.4 Nutritional Status

1.5 Challenges for Genetic Enhancement

1.6 Characteristics and Cultivars

References

Chapter 2. Taxonomy and Origin

2.1 Taxonomy

2.2 Origin

References

Chapter 3. Morphology and Breeding Behavior

3.1 Morphology

3.2 Breeding Behavior and Pollination Control

References

Chapter 4. Genetic Variability for Qualitative and Quantitative Traits

4.1 Morphological/Phenotypic Level

4.2 Biochemical Level

4.3 DNA Level

References

Chapter 5. Genetics and Cytogenetics

5.1 Genetics

5.2 Cytogenetics

References

Chapter 6. History of Winter Sorghum Improvement in India

6.1 The Origin of M 35-1

References

Chapter 7. Breeding Methods for Winter Sorghum Improvement

7.1 Yield and Adaptation Breeding for Grain and Fodder Yield

7.2 Breeding for Resistance to Abiotic Stresses

7.3 Breeding For Resistance To Diseases

7.4 Breeding For Resistance To Insect Pests

7.5 Breeding for Grain, Fodder, and Nutritional Quality

7.6 Participatory Varietal Selection

References

Chapter 8. Industrial or Alternate Uses

8.1 Industrial Products

References

Index

Copyright

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Preface

Sorghum (Sorghum bicolor (L.) Moench) is the fifth important cereal crop in the world, primarily grown for grain production on about 35 m ha in about 106 countries and a staple food crop for over 500 million people in Africa, Asia, Oceania, and the Americas. The crop is mainly grown in tropical and subtropical areas which are marginal and stress prone. The postrainy sorghums or the winter sorghums are specialized sorghums of India. They are agronomically and physiologically distinct from the rainy sorghums and adapt well to the drought conditions as they have been grown and selected under receding moisture conditions occurring after cessation of rains. Unlike rainy sorghum where the grain quality is affected due to grain mold infection, the winter sorghum/postrainy sorghum is the main source of food as it is free from grain mold. It also serves as an important source of fodder since fodder from other crops is not available during this season. As not much progress has been made in the improvement of the postrainy sorghums, the landraces with tolerance to shoot fly, terminal drought and charcoal rot and with bold, round and lustrous grain dominate the winter sorghum tracts. Rapid productivity enhancement has not been possible for winter sorghum unlike rainy season grown sorghum due to lack of success with hybrid technology. However winter sorghum is an important crop in lieu of climate change and also as a health crop. Though several researchers worked for winter sorghum improvement in Indian Council of agricultural research (ICAR), India and International Crops Research Institute for the Semi Arid Tropics (ICRISAT), the information is scattered and not compiled. Hence, an attempt is made to pool the available literature to give a glimpse on what has gone through in decades of research on winter sorghum improvement.

This book has eight chapters. The first five chapters deal with crop biology, origin and taxonomy, genetic variability, genetics and cytogenetics. These do not tell about actual breeding, but a knowledge about these is useful in enhancing the effectiveness of a breeding programme. The sixth chapter deals with history of winter sorghum improvement in India. Winter sorghum research did not receive much emphasis until nineties and the varieties or hybrids bred and released could not match M 35-1, a landrace variety that is grown for about seven and a half decade, in yield or quality. However, lot of strategic research is required to develop new varieties and hybrids for post-rainy season adaptation that can break the yield plateau. The history of past research for winter sorghum improvement will help to plan future strategies to bring in real impact at the farmers level. The seventh chapter on conventional and molecular breeding for sorghum improvement deals with strategies for breeding for yield and adaptation, tolerance to abiotic stresses influencing winter sorghum (terminal drought and mid-season cold stress) and tolerance to biotic stresses limiting yields of winter sorghum (charcoal rot among diseases and shootfly, aphids and shoot bug among insects). The eighth chapter exposes the reader on alternate and industrial uses of sorghum.

Sorghum’s place in most of its traditional environments will remain important, with great potential in non-traditional environments. We hope that the "Genetic Enhancement of rabi sorghum: Adapting the Indian Durras" will make a valuable reference book for students, teachers, researchers interested in winter sorghum research and development.

Editors

Chapter 1

Introduction

The chapter gives an introduction to sorghum in general, and winter sorghum in particular. Importance of postrainy/winter sorghum is discussed in terms of production and uses. The general aspects of winter sorghum, its nutritional advantages, climatic requirements, characteristics, cultivars, and challenges for genetic enhancement are briefly discussed.

Keywords

Winter sorghum; food; fodder; gluten free; photoperiod sensitive; terminal moisture stress

Outline

1.1 Production Statistics 2

1.2 Environmental Factors Limiting Rabi Sorghum Productivity 4

1.2.1 Climatic Factors 4

1.2.2 Edaphic Factors 4

1.3 Uses 5

1.4 Nutritional Status 6

1.5 Challenges for Genetic Enhancement 6

1.6 Characteristics and Cultivars 6

References 7

Sorghum [Sorghum bicolor (L.) Moench] is cultivated as a major food crop in several countries in South Asia, Africa, and Central America. The crop is mainly grown in tropical and subtropical areas that are marginal and stress prone. Sorghum is known as guinea-corn, dawa, or sorgho in West Africa, durra in the Sudan, mshelia in Ethiopia and Eritrea, mtama in East Africa, kaffircorn, mabele, or amabele in southern Africa, and as jowar, jonna, cholam, or jola in the Indian subcontinent (Bantilan et al., 2004).

Sorghum is grown mainly for animal feed in the United States, Australia, and South America, while it is mainly grown as food in Africa and India. In India, sorghum is grown in two seasons, the kharif (or rainy) season and the rabi (or postrainy) season. During rabi, it is sown between September and end of October in the Deccan Plateau between 10° and 20°N latitude (Seetharama et al., 1990), and it accounts for 45% of the total sorghum area under cultivation and 32% of the total production (Sajjanar et al., 2011). In India, sorghum grown in the rainy season is mainly utilized as feed, as the grain is often caught in rains prevailing during harvesting and the grain quality is affected due to molds. However, postrainy sorghum is primarily used as a food owing to its good grain quality, and it also serves as a main source of fodder, especially during dry seasons, and is an important crop in lieu of food security. Most of the rabi sorghum varieties belong to durra or intermediates of the durra race, while the kharif cultivars that are being grown belong to caudatum and kafir races (Reddy et al., 2003). The majority of rabi germplasm samples, which mostly belonged to the durra race, were collected by the National Bureau of Plant Genetic Resources (NBPGR) and International Crops Research Institute for the Semi-Arid Tropics (ICRISAT; Mathur et al., 1993). From the breeding point of view, kafir, caudatum, and durra, having genes contributing to yield, have been extensively utilized in breeding programs across the globe. Despite some major differences in both environment and cultural practices, rabi sorghum grown in India is similar to sorghum grown on residual moisture elsewhere in Africa. However, two major differences exist: African postrainy sorghums are grown in low soil fertility conditions, as they are cropped on receding flood plains after burning vegetation; and low plant density is employed (Seetharama et al., 1990) (Figure 1.1).

Figure 1.1 Rabi sorghum panicle.

1.1 Production Statistics

Sorghum [S. bicolor (L.) Moench] is the fifth most important crop in acreage next only to wheat, rice, maize, and barley in the world (Dillon et al., 2007). It was cultivated on 35 m ha in 106 countries in 2011 and a staple food crop for over 500 million people in Africa, Asia, Oceania, and the Americas. In 2011, India was the world’s largest producer of sorghum (7.0 million metric tons), followed by Nigeria (6.9), Mexico (6.4), the United States (5.4), and Argentina (4.5). India is also the largest cultivator of sorghum (7.4 m ha), followed by Nigeria (4.9), Niger (2.9), and Ethiopia (2.2) (FAOSTAT, 2013).

Source: FAOSTAT (2013).

During the last 30 years (1981–2011), the annual world production of sorghum has decreased from 73.3 to 54.2 million tons, and the area planted decreased from 45.9 to 35.5 million ha. However, the average yields in 1981 and 2011 were almost similar (1598 and 1527 kg/ha). However, wide variations exist between these global figures and figures at the national level. In India, for example, between 1981 and 2011, the area planted to sorghum fell from 16.6 to 7.4 million ha (Figure 1.2), and annual production fell from 12.1 to 7.0 million tons (Figure 1.3), but yields increased by 31% from 727 (in 1981) to 949 kg/ha (in 2011) (Figure 1.4). There exists a wide gap between average yield levels of India compared to the world yield levels.

Figure 1.2 Trends in area under sorghum for three decades.

Figure 1.3 Trends in sorghum production for three decades.

Figure 1.4 Trends in sorghum productivity for three decades.

Sorghum is well adapted to hot and dry (semiarid) agro-ecologies, where it is difficult to grow other food grains. Since sorghum is grown in drought-prone areas, it is cultivated with minimal inputs; in conditions of low soil fertility; under rainfed conditions with erratic and inadequate rainfall using traditional cultivars, which are poor yielders; and also exposed to a variety of disease and pest problems that affect the crop yields (Rai et al., 1999). The lower average yields are primarily due to the hot, dry conditions where sorghum is mostly grown, rather than the plant’s own capability. However, sorghum has high yield potential, comparable to rice, wheat, and maize (House, 1985). Where moisture is not a limiting factor, sorghum had yielded up to 11,000 kg/ha at the field level, with average yields ranging from 7000 to 9000 kg/ha. In traditional sorghum-cultivating areas, yields of 3000–4000 kg/ha are obtained under better management conditions, dropping to 300–1000 kg/ha when moisture and soil fertility become limiting factors (House, 1985).

1.2 Environmental Factors Limiting Rabi Sorghum Productivity

1.2.1 Climatic Factors

Rainfall: The rabi crop requires about 175 mm of water (Tarhalkar, 1986). Most rabi sorghum is grown in areas receiving annual rainfall of about 600–800 mm. The probability of receiving 10 mm of rainfall during the first week of October is about 60%, and soon after, it decreases to about 2–5% per week (Virmani et al., 1982).

Solar radiation: The mean daily radiation is similar during both kharif and rabi seasons, being only 6% less in rabi season (Sivakumar and Virmani, 1982). However, the conversion of incident solar radiation to dry matter is only half in rabi season, as opposed to kharif season (Sivakumar and Huda, 1985), due to the lower leaf area during rabi and low radiation use efficiency (Seetharama et al., 1982).

Temperature: The mean temperature during rabi season (24.9°C) was reported to be only marginally less than kharif season (27.9°C) at Patancheru, Hyderabad (Sivakumar and Virmani, 1982). However, the variations in the temperature during the day are greater in rabi than kharif season, and the low night temperatures are believed to be a main cause for the reduction in growth and grain yield (Rao et al., 1977; Choudhari, 1989). Minimum temperatures decline from about 20°C at sowing to 12°C at flowering, and increase to 18°C during grain-filling period (Mukri et al., 2010). Early-sown crop suffers from cold temperature stress during the flowering stage, resulting in poor seed sets, especially in hybrids. Late-sown crop suffers from cold temperature stress at the vegetative stage, resulting in profuse tillering. The late-sown crop also suffers from high temperature stress, coupled with low soil moisture and high evaporative demand at the grain-filling stage.

Photoperiod: The change in photoperiods at panicle initiation across different sowing dates is less in rabi sorghum, but its interaction with temperature may be of considerable significance (Seetharama et al., 1990).

1.2.2 Edaphic Factors

Rabi sorghum is grown in varied depths of soil. Much of it is grown in black soils capable of holding moisture. The soils show a great variability in their depth and characteristics and are grouped as shallow, medium, and deep black soils. The depth of shallow soils varies from 0.0 to 22.5 cm, and about 20–22% of the black soils are classed as shallow soils. The soil depth of medium soils is about 22.5–90 cm. About 65% of the area is comprised of medium black soils. Deep black soils are more fertile than medium black soils, have a granular structure in the surface layer, and become cloddy at lower depths. The clay content of black soils ranges from 40% to 60% but may be as high as 70%. The differences in grain yield between shallow and deep vertisols is around 1.0 t/ha (Tandon and Kanwar, 1984).

1.3 Uses

Sorghum grain is mostly consumed directly for food (55%) in the form of porridge (thick or thin) and flatbread. It is also an important source of feed grain (33%), especially in Australia and the Americas. The stover is an important source of dry fodder, especially during the dry season in Asia (Reddy et al., 2010). Sorghum has great potential as a fodder resource due to its quick and rapid growth, high green fodder yield, and good quality. Of late, sweet sorghum is emerging as an important biofuel crop, making sorghum a unique crop with multiple advantages as food, feed, fodder, fuel, and fiber. Hence, it is popularly known as a smart crop. In addition to these uses, sorghum crop residues and green plants provide building material, and fuel for cooking, particularly in the dry land areas (Chandel and Paroda, 2000), and in paper and cardboard, jaggery, and ethanol production. Sorghum has great potential to provide raw material for industries. Industrial application of sorghum makes its cultivation economically viable for marginal farmers. The grain is used for industrial purposes, such as potable alcohol, malt, beer, liquids, gruels, starch, adhesives, core binders for metal casting, ore refining, and grits as packaging material (Reddy et al., 2006) (Figure 1.5).

Figure 1.5 Uses of sorghum.

1.4 Nutritional Status

Sorghum grain contains 8–15% protein, 5–15% sugar, and 32–57% starch. It is relatively rich in micronutrients (mg/kg) iron (41–127), zinc (14–35), phosphorus (1498–3797), Ca (207–447), K (1150–2569), Mn (10–24), Na (12–54), and Mg (750–1506) (Shegro et al., 2012). Tannins, found in red-grained sorghums, contain antioxidants that protect against cell damage, a major cause of disease and aging. The protein and starch in sorghum grain are comparatively slowly digested than other cereals. This aspect is highly beneficial for people with diabetes; hence, sorghum is considered health food. Sorghum is a good alternative to wheat flour for individuals suffering from celiac disease, as the starch from sorghum grain is gluten free.

1.5 Challenges for Genetic Enhancement

Sorghum is cultivated by poor farmers and grown under subsistence conditions. Hence, they cannot take advantage of high yield potential, as the growers are unable to follow improved management practices. Higher yields can be obtained by growing varieties/hybrids with improved tolerance to drought, heat, and low soil fertility, as well as resistance to pests and diseases. Pest problems comprise one of the major constraints for achieving higher yields in sorghum grown in tropical areas. Immense losses are caused by insect pests attacking sorghum at all stages of growth, the important ones being shoot fly and aphids in winter sorghum. Pests like the shoot fly and stem borer attack in the very early stages of sorghum, resulting in the complete loss of the crop. Aphids attack from the flowering to milk stages and cause both qualitative and quantitative yield losses. The midge and ear-head bugs attack at the grain-filling stage leading to the losses up to 100%.

1.6 Characteristics and Cultivars

Winter sorghums are characterized by their response to shorter day lengths (photoperiod sensitivity), flowering, and maturity (occurring more or less at the same time irrespective of temperature fluctuations) and sowing dates (thermo-insensitivity within the postrainy season varieties). They are tolerant to terminal moisture stress and resistant to stalk rot/charcoal rot. As the fodder is as important as grain, the varieties are selected to produce high biomass (grain and stover) and have high lustrous grain with semi-corneous endosperm. Tolerance to shoot fly, lodging (mechanical), and rust are also required (Sanjana Reddy et al., 2012). All these characters exist in M 35-1, a variety selected from a local landrace nearly 75 years ago at Mohol in Maharashtra, which produces high stable yields of grain and stover across different sowing dates. As a result, M 35-1, a landrace selection developed in 1937 still dominates the postrainy season sorghum areas in India (Sanjana Reddy et al., 2009). In the past, focused breeding efforts on rabi sorghum led to the development of several rabi sorghum varieties such as CSV8R, Swati, CSV14R, CSV 18R, CSV216R, and CSV22R. Heterosis breeding led to the release of hybrids like CSH8R, CSH12R, CSH13R, CSH15R, and CSH19R. However, these varieties have become more popular compared to hybrids, a situation quite opposite to kharif. Though the hybrids are heterotic for grain yield, they have poor grain quality and are vulnerable to biotic and abiotic stresses (Shinde et al., 2010).

References

1. Bantilan MCS, Deb UK, Gowda CLL, Reddy BVS, Obilana AB, Evenson RE. Introduction. In: Bantilan MCS, Deb UK, Gowda CLL, Reddy BVS, Obilana AB, Evenson RE, eds. Sorghum Genetic Enhancement: Research Process, Dissemination and Impacts. Patancheru, Andhra Pradesh, India: International Crops Research Institute for the Semi-Arid Tropics; 2004:5–18.

2. Chandel KPS, Paroda RS. Status of plant genetic resources conservation and utilization in Asia-Pacific region – Regional synthesis report Asia-Pacific Association of Agricultural Research Institutions Bangkok, Thailand: FAO Regional Office for Asia and the Pacific; 2000; 158 p.

3. Choudhari, S.D., 1989. Production technology and a new line of work to improve Rabi sorghum production. Paper Presented During the Working Group Meeting on Production Technology of Rabi Sorghum. 19 September 1989, CRIDA, Hyderabad.

4. Dillon SL, Shapter FM, Henry RJ, Cordeiro G, Izquierdo L, Slade LL. Domestication to crop improvement: genetic resources for sorghum and saccharum (Andropogoneae). Ann Bot (Lond.). 2007;100:975–989.

5. FAOSTAT, 2013. Agricultural data. Available from: <http://apps.fao.org> (accessed May 2013).

6. House LR. A Guide to Sorghum Breeding second ed. Patancheru, India: International Crops Research Institute for the Semi-Arid Tropics; 1985.

7. Mathur PN, Gopal Reddy V, Prasada Rao KE, Mengesha MH. Collection of rabi sorghum germplasm I Northern Karnataka and adjoining areas of Andhra Pradesh. Indian J Plant Genet Resour. 1993;6:1–8.

8. Mukri G, Biradar BD, Sajjanar GM. Effect of temperature on seed setting behavior in rabi sorghum (Sorghum bicolor (L.) Moench). Electron J Plant Breed. 2010;1(4):776–782.

9. Rai KN, Murty DS, Andrews DJ, Bramel-Cox PJ. Genetic enhancement of pearl millet and sorghum for the semi-arid tropics of Asia and Africa. Genome. 1999;42:617–628.

10. Rao, N.G.P., Vidyabhushanam, R.V., Rana B.S., 1977. Recent development in sorghum breeding in India; Pages 13 to 18 in Section 7, Plant Breeding Papers, Third International Congress of the Society for the Advancement of Breeding Researches in Asia and Oceania (SABRAO), February, 1977, Canberra, Australia.

11. Reddy, B.V.S., Sanjana, P., Ramaiah, B., 2003. Strategies for improving post-rainy season sorghum: a case study for landrace hybrid breeding approach. Paper presented in the Workshop on Heterosis in Guinea Sorghum, Sotuba, Mali, pp. 10–14.

12. Reddy BVS, Ramesh S, Reddy PS. Sorghum genetic resources, cytogenetics, and improvement. In: Singh RJ, Jauhar PP, eds. Genetic Resources Chromosome Engineering and Crop Improvement, Cereals, vol 2. Boca Raton, FL: CRC Press, Taylor & Francis Group; 2006:309–363.

13. Reddy BVS, Ashok Kumar A, Sanjana Reddy P. Recent advances in sorghum improvement research at ICRISAT. Kasetsart J (Natural Science). 2010;44:499–506.

14. Sajjanar GM, Biradar BD, Biradar SS. Evaluation of crosses involving rabi landraces of sorghum for productivity traits. Karnataka J Agric Sci. 2011;24(2):227–229.

15. Sanjana Reddy P, Reddy BVS, Ashok Kumar A. M 35-1 derived sorghum varieties for cultivation during the postrainy season. E-J SAT Agric Res. 2009;7.

16. Sanjana Reddy P, Patil JV, Nirmal SV, Gadakh SR. Improving post-rainy season sorghum productivity in medium soils: does ideotype breeding hold a clue? Curr Sci. 2012;102:904–908.

17. Seetharama N, Reddy BVS, Peacock JK, Bidinger FR. Sorghum improvement for drought resistance. In: Drought Resistance in Crop Plants with Emphasis on Rice. Philippines: IRRI; 1982:317–356.

18. Seetharama N, Singh S, Reddy BVS. Strategies for improving postrainy sorghum productivity. Proc Indian Natl Sci Acad. 1990;56(5&6):455–467.

19. Shegro A, Shargie NG, van Biljon A, Labuschagne MT. Diversity in starch, protein and mineral composition of sorghum landrace accessions from Ethiopia. J Crop Sci Biotechnol. 2012;15(4):275–280.

20. Shinde DG, Biradar BD, Salimath PM, Kamatar MY, Hundekar AR, Deshpande SK. Studies on genetic variability among the derived lines of B×B, B×R and R×R crosses for yield attributing traits in rabi sorghum (Sorghum bicolor (L.) Moench). Electron J Plant Breed. 2010;1(4):695–705.

21. Sivakumar MVK, Huda AKS. Solar energy utilization by tropical sorghums. Agric Forest Meteorol. 1985;35:47–57.

22. Sivakumar MVK, Virmani SM. The physical environment; in Sorghum in the Eighties. In: International Crops Research Institute for the Semi-Arid Tropics, Patancheru, India. 1982:83–100.

23. Tandon, H.L.S., Kanwar, J.S., 1984. A Review of Fertilizer Use Research on Sorghum in India. Research Bulletin No. 8. International Crops Research Institute for the Semi-Arid Tropics, Patancheru, A.P. India.

24. Tarhalkar, P.P., 1986. Agronomical investigations on rabi sorghum: a brief review. Presented at the AICSIP Annual Workshop. 14–16 May 1986, Andhra Pradesh Agricultural University, Hyderabad.

25. Virmani SM, Sivakumar MVK, Reddy SJ. Rainfall probability estimates for selected locations of semi-arid India Patancheru, Andhra Pradesh, India: International Crops Research Institute for the Semi-Arid Tropics (ICRISAT); 1982; (Research Bulletin No. 1), 170 pp.

Chapter 2

Taxonomy and Origin

Most winter sorghums grown in India belong to the race known as durra. The taxonomy of cultivated sorghum and the classification of races are presented. The origin of durras and their transformation into specialized sorghums grown in winter season is explained.

Keywords

Race; durra; origin; taxonomy

Outline

2.1 Taxonomy 9

2.2 Origin 11

References 13

2.1 Taxonomy

Sorghum was first described by Linnaeus in 1753 under the name Holcus. In 1794, Moench distinguished the genus Sorghum from genus Holcus (Celarier, 1959; Clayton, 1961) and brought all the sorghums together under the name Sorghum bicolor (House, 1978; Clayton, 1961). Subsequently, several authors have discussed the systematics, origin, and evolution of sorghum since Linnaeus (de Wet and Huckabay, 1967; de Wet and Harlan, 1971; Doggett, 1988). Dahlberg (2000) provides an excellent overview of the present-day classification to describe the variation found within cultivated sorghums.

Sorghum is classified under the family Poaceae, tribe Andropogoneae, subtribe Sorghinae, and genus Sorghum Moench (Clayton and Renvoize, 1986). Garber (1950) and Celarier (1959) further divided the genera into five subgenera: sorghum, chaetosorghum, heterosorghum, parasorghum, and stiposorghum. S. bicolor was further broken down into three subspecies: S. bicolor subsp. bicolor, S. bicolor subsp. drummondii, and S. bicolor subsp. verticilliflorum. The cereal sorghums were found to consist of four wild races and five cultivated races (Harlan et al., 1976). The four wild races of S. bicolor are arundinaceum, virgatum, aethiopicum, and verticilliflorum. They are placed in S. bicolor subsp. verticilliflorum, formerly subsp. arundinaceum. Cultivated sorghums are classified as S. bicolor subsp. bicolor and are represented by several agronomic types, such as grain sorghum, sweet sorghum, sudangrass, and broomcorn (Berenji and Dahlberg, 2004). Additionally, there are two weedy sorghums widespread in temperate zone; that is, Johnsongrass and spontaneous sorghum (shattercane).

S. bicolor subsp. bicolor contains cultivated sorghum races. The cultivated races that are presently conceived are bicolor, guinea, kafir, caudatum, and durra. Intermediates that are caused by hybridization of these races exhibit characters of both parents (Smith and Frederiksen, 2000). Harlan and de Wet (1972) classified S. bicolor (L.) Moench, subsp. bicolor into 5 basic and 10 hybrid races as depicted next.

The 15 races of cultivated sorghum are identified by mature spikelets, and this classification is based on five fundamental spikelet types (Harlan and de Wet, 1972). The International Plant Genetic Resources Institute Advisory Committee on Sorghum and Millets Germplasm has accepted and recommended this classification for describing sorghum germplasm (IBPGR and ICRISAT, 1993). These races are known to differ significantly not only for grain quality traits, but also for yield potential. The interracial hybrids were found to have greater heterosis than those of intraracial hybrids. Knowledge of the racial characteristics helps the breeder achieve systematic genetic improvement of sorghum for traits of interest (Reddy et al., 2008) (Figure 2.1).

Source: Reddy et al. (2002).

Figure 2.1 Races of S. bicolor.

2.2 Origin

Mann et al. (1983) hypothesized that the origin and early domestication of sorghum took place approximately 5000 years ago in northeastern Africa. Wendorf et al. (1992) reported new evidence that places the origin and domestication at 8000 years before present (BP) on the Egypt-Sudan border. Thus, there seems to be no argument against the African origin of sorghum (Kimber, 2000), a concept that is also supported by the largest diversity of the cultivated and wild sorghum in Africa (de Wet, 1977; Doggett, 1988). The great diversity of S. bicolor has been created through disruptive selection (i.e., selection for extreme types) and by isolation and recombination in the extremely varied habitats of northeast Africa and the movement of people carrying the species throughout the continent (Doggett, 1988). On the Indian subcontinent, evidence for early cereal cultivation was discovered at an archaeological site in the western part of Rojdi (Saurashtra) dating to about 4500 BP (Damania, 2002). The Indian subcontinent is considered to be the secondary center of origin of sorghum (Vavilov, 1992).

Bicolor is widely distributed in Africa and coastwise from India to Indonesia and then to China (de Wet and Price, 1976). It appears that this race arose in east Africa from the subsp. aethiopicum, and the great diversity found in Asia occurred after its introduction there. However, according to Dahlberg (1995), early bicolors are believed to have arisen from the subspecies verticilliflorum in central Africa, and they were thought to have introgressed with wild forms and gave rise to the races caudatum, kafir, guinea, and durra (Dahlberg, 1995). As the bicolors moved to the west, they came into contact with wild Sorghum arundinaceum, from which the race guinea evolved (Dahlberg, 2000). Harlan and Stemler (1976) considered guineas to be the oldest of the races, mostly found in western and eastern Africa. The morphological affinities and distribution indicate that the guinea race was probably derived from selection among wild members of the subsp. arundinaceum. The caudatum race is thought to have been derived from an introgressed cross of an early bicolor with a wild sorghum and arose from the area of early bicolor domestication (Dahlberg, 2000). According to Stemler et al. (1975), the caudatum race was domesticated later than bicolor and guinea and thought to be segregated out of bicolor. It is dominant in parts of Sudan, Chad, Nigeria, and most parts of Uganda. This is an important race agronomically, especially in combination with other races. The bicolors crossed with wild verticilliflorum in northern Africa and gave rise to the kafir race and carried by the Bantu speakers of Africa to the east and south (Dahlberg, 1995). According to de Wet (1978) and Harlan et al. (1976), the kafir race was derived from an early bicolor race and may have migrated before the guinea race was segregated from the bicolor race (Smith and Frederiksen, 2000). It is cultivated from northern Nigeria to west to northern Ghana, where there is a gene flow between the guinea and kafir races. Its distribution and morphological affinities suggest that it arose from the subsp. verticilliflorum. The durra race is thought to have originated in Ethiopia from early bicolors, which introgressed with wild aethiopicum adapted to drier conditions (Dahlberg, 1995). Doggett (1988) also argued that the durra race is Ethiopian in origin and its introgression with wild forms permitted adaptation to drier conditions that developed in the highlands. These upland races descended to the lowlands, and the adapted varieties migrated west through Yemen and Saudi Arabia to India.

The durra race has a compact panicle, indicating adaptation to low-rainfall environment (Mann et al., 1983). Durras are distributed in a belt of 10–15° N latitude from Ethiopia to Mauritania; that is, in the mid-altitude highlands of Ethiopia, the Nile Valley of Sudan and Egypt, and in India and Pakistan (Kimber et al., 2013). There appears to be three centers of morphological diversity: the Ethiopian-Sudan region, the near East, and India (Smith and Frederiksen, 2000).

Most of the winter sorghums grown in India belong to the durra race. Harlan and Stemler (1976) felt that durra sorghums were selected from early bicolor, which had been carried to India before 3000 BP and may have been domesticated in India. Doggett (1988) argues that durras originated in Ethiopia since the whole sequence of wild type-bicolor-durra is clearly represented there. Doggett (1988) suggested that the early bicolors introgressed with wild forms, and they got adapted to drier conditions, which led to the development of the durra race. The durras moved to the west through Sudan and began to occupy the drier regions below the southern margin of the Sahara. Then they moved through the Horn of Africa and worked their way to India. The compact panicle and predominantly white seeds of the durra race are indications that it adapted to low-rainfall environments with a low risk of grain mold (Mann et al., 1983). However, white-grain sorghum is desirable as human food because of its low tannin content and plant improvement programs continue to expand its area of adaptability (Smith and Frederiksen, 2000). The durras were widely cultivated by Muslim Africans and Arabic people in Ethiopia. In Ethiopia, the Muslim Oromos (Gallo), who settled in the fertile warm highland almost 500 years ago cultivated the durra race, which formed the foundation of their agricultural system (Harlan et al., 1973). The durras are presently distributed in the mid-altitude highlands of Ethiopia, the Nile Valley of Sudan and Egypt, and in a belt 10–15° N latitude from Ethiopia to Mauritania. They also are grown in the Islamic and Hindu areas of India and Pakistan (Smith and Frederiksen, 2000). Haaland (1995, 1998) proposed a hypothesis called the Haaland hypothesis. The Haaland hypothesis proposes that wild S. bicolor was exported to India, where it became domesticated durra and was subsequently reintroduced into Africa during Islamic times. In India, durra reaches its most extreme forms with creases on both glumes, while in Africa, it is often modified through hybridization with other races.

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