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Utilization of Wild Relatives of Pearl Millet W . W . H a n n a 1 Abstract The wild Pennisetum species offer diverse germplasm that can be used to improve pearl millet. In the primary gene pool, the wild, weedy pearl millet subspecies monodii and stenostachyum are the most readily and easily utilized. The germplasm in the secondary and tertiary gene pools is more difficult to manipulate and transfer to pearl millet. Useful characteristics in the wild species that could be used to improve pearl millet include disease and insect resistance, genes for fertility restoration of the A 1 cytoplasm, cytoplasmic diversity, yield genes, apomixis, maturity, and many inflorescence and plant morphological characteristics. Research with the wild species requires specific objectives, large populations, good screening methods, and a team effort. One of the greatest research needs in the Pennisetum genus is adequate and systematic collection and preservation of the wild species. Introduction Pearl millet (Pennisetum americanum) is an excellent research organism, an important world food and forage crop, and has abundant natural genetic diversity, but it has not received the attention it deserves. Its potential contribution to science as a research organism and its potential for meeting world food needs as a grain crop have not been fully recognized. Pearl millet makes an excellent research organism because of many desirable characteristics. It is a diploid with 2n = 14 large chromosomes, and has a protogynous habit of flowering which allows it to be easily cross-pollinated and genetically manipulated. A single plant can be readily selfed and crosspollinated as both male and female. Inbreds are usually vigorous and hybrid vigor is significant. An inflorescence can produce 1000 or more seeds and a single-spaced plant can produce 25 or more inflorescences. Some genotypes will naturally flower 35-40 d after planting, or can be induced to do so with short day length and high temperatures, thus 1. Research Geneticist. USDA-ARS. Coastal Plain Experiment Station, Department of Agronomy, Tifton, Georgia 31793, USA. ICRISAT (Internationa! Crops Research Institute for the Semi-Arid Tropics). 1987. Proceedings of the International Pearl Millet Workshop, 7-11 April 1986. ICRISAT Center, India. Patancheru, A.P. 502324, India: ICRISAT. 33
making it possible to grow a number of generations per year. Pearl millet plants can grow to maturity in a 5-cm pot and produce 100 or more seeds. In the opinion of the author, these and other characteristics make pearl millet the "Drosophila" of the plant kingdom. Pearl millet is an important food grain in Africa and India. It is a high quality forage in the United States, Australia, and South America. However, there is new interest in growing pearl millet for grain in the USA because it is drought tolerant and produces high quality grain. Pearl millet tolerates low fertility and low soil pH, but responds positively to more favorable soil and water conditions. Evolutionary History and Domestication of Pearl Millet Based on evidence of genetic diversity within the genus in Africa, it is generally accepted that pearl millet originated there. More specifically, the Sahelian Zone from western Sudan to Senegal appears to be the center of origin (Harlan 1975). Various reports indicate pearl millet has been grown for thousands of years (Brunken 1977, Burton and Powell 1968, Rachie and Majmudar 1980). In the last 200-300 years, pearl millet has been given a number of genera and species names such as P. glaucum L. and P. typhoides (Burm.) Stapf and Hubb. In 1976 it was renamed P. americanum (L.) Leeke (see Terrell 1976). Those interested in more detailed discussions on the taxonomy, origin, domestication, and evolutionary history of pearl millet should consult Brunken (1977), Jauhar (1981), Rachie and Majmudar (1980), and Stapf and Hubbard (1934). Gene Pools Pennisetum is a genus with over 140 species (Brunken 1977) divided into five sections: Gymnothrix, Eu- Pennisetum, Penicillaria, Heterostachya, and BrevivaJvuia (Stapf and Hubbard 1934). Pearl millet belongs to the Pcnicillaria section. The literature and observations by the author at the Royal Botanic Gardens, Kew, U K , indicate that most of the wild Pennisetum species are found in Africa. However, there are a number of reports of Pennisetum species being collected in other countries. There are specimens of Pennisetum species on record at Kew from Israel, Japan, India, Australia, New Zealand, Polynesia, Mexico, Central and South America, and other countries, including many in Africa. The Pennisetum genus has species with chromosome numbers in multiples of x = 5, 7, 8, and 9 (see Table 1, Hanna and Dujardin In press, and Jauhar 1981). In the x = 5 group is P. ramosum (2n = 10). The x = 7 group includes pearl millet and its wild weedy subspecies (2n = 14), P. purpureum (2n = 28), and P. schweinfurthiii (2n = 14). Pennisetum massaicum (2n = 16 and 32) is the only known species in the x - 8 group. The remainder of the species where chromosome numbers have been established belong to the x = 9 group. Ploidies reported range from diploid to octoploid (Jauhar 1981). Both sexual and apomictic species as well as annual and perennial species are included in the genus. The diverse germplasm in Pennisetum, both within and between species, offers possibilities for use in improving pearl millet and to produce interspecific hybrids with forage potential. The ease with which the wild germplasm can be manipulated and used will vary both within and between species. One way of classifying the wild or exotic germplasm is by the ease with which it can be introgressed into the cultivated species. A system of primary, secondary, and tertiary gene pools has been suggested for dividing the wild germplasm (Harlan and de Wet 1971). In Pennisetum, the primary gene pool includes the wild, grassy, and weedy P. americanum subspecies (2n = 14), monodii and stenostachyum that readily cross with pearl millet and produce fertile hybrids (Table 1). Monodii is the true weedy subspecies found mainly in the Sahelian Zone of Africa in an east-west line from Senegal to Ethiopia. It is found "as a natural colonizer on sandy soils in disturbed habitats such as seasonal stream beds and roadsides as well as a weed near human habitations'* (Brunken 1977). Inflorescences are 2-20 cm long with loosely arranged involucres that readily shatter (Brunken 1977). Inflorescence length in about 100 accessions that the author observed ranged from 2-12 cm. Involucral bristles are densely plumose. Stems are thinner and leaves are narrower than those of most pearl millet accessions. Most flower in response to short day length. Monodiiis a rich source of germplasm that can be used to improve pearl millet. Data indicate that it is an excellent source of genetic diversity for new cytoplasms, stable cytoplasmic sterility, pest resistance (disease and insect), fertility restoration, hybrid vigor, etc. Based on intuition and five years of research with monodii, the author believes that this 34
Page 2 and 3: Cover: Ex-Bornu, an important landr
Page 4 and 5: Correct citation: I C R I S A T (In
Page 6 and 7: Pearl Millet Entomology Insect Pest
Page 8 and 9: Potential and Prospects of Pearl Mi
Page 10 and 11: tropical countries i t w i l l be v
Page 12: Opening Session
Page 15 and 16: globusum has globose caryopses, and
Page 17 and 18: Area, Production, and Productivity:
Page 19 and 20: area increased substantially in Raj
Page 21 and 22: apply complex fertilizer at 20-60 k
Page 23 and 24: Diseases. Diseases are endemic to p
Page 25 and 26: levels of adoption. The relative co
Page 27 and 28: Socioeconomic Factors Management. T
Page 30 and 31: Pearl Millet in African Agriculture
Page 32 and 33: The poor performance in food produc
Page 34 and 35: 900 800 700 600 Mean 500 400 300 20
Page 36 and 37: population pressures in recent year
Page 38 and 39: Economics of Millet Production In 1
Page 40 and 41: ather than to yield increases, show
Page 42: Simpara, M. B. 1985. La recherche s
Page 47 and 48: in about 30 accessions. Unlike mono
Page 49 and 50: genome male-sterile lines, and A A
Page 51 and 52: Pearl m i l l e t (Pennisetum ameri
Page 53 and 54: Vasil, V., and Vasil, I . K . 1981a
Page 55 and 56: Introduction Pearl millet (Penniset
Page 57 and 58: Clean sorghum o r m i l l e t G r i
Page 60 and 61: peas, peanuts, or baobab leaves. Wh
Page 62 and 63: Beer Two major kinds of beer are pr
Page 64 and 65: properties for pearl millets. It is
Page 66 and 67: 54 Figure 9. A. Pearl millet with a
Page 68 and 69: the germ (McDonough 1986). The high
Page 70 and 71: Table 6. Nitrogen distribution in s
Page 72 and 73: Banigo, E.O.I., de Man, J.B., and D
Page 75 and 76: Discussion The discussion of the pa
Page 77: Improvement through Plant Breeding
Page 81 and 82: Diversity and Utilization of Pearl
Page 83 and 84: with protruding grains, but in the
Page 85 and 86: Table 2. Pearl millet accessions as
Page 87 and 88: lese millets constitute two distinc
Page 89 and 90: turn, P. purpureum, (Dujardin and H
Page 91 and 92: more to useful genetic diversificat
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Hanna, W . W . , Wells, H . D . , a
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Varietal Improvement of Pearl Mille
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the weedy form can significantly in
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Table 3. Evaluation of heterosis in
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Table 4. Grain yield of local 3/4 p
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ased on selection of drought-resist
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Marchais, L., and Pernes, J. 1985.
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Andrews 1984), there are operationa
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S h o r t - t e r m g o a l s I n t
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Table 4a. Prediction equations, adv
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Table 4c. Prediction equations, adv
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Nebraska B s y n t h e t i c o r i
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6 P o p u l a t i o n c r o s s e s
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Pearl Millet Hybrids H . R . Dave 1
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Table 2. Early released hybrids in
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Table 6. Performance of third phase
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Breeding Pearl Millet Male-Sterile
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Table 1. Male-sterile lines of pear
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Senegal. This source is reported to
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possible solutions to produce high
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selected from IP 2696 has recently
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Burton, G.W., and Athwal, D.S. 1967
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Biological Factors Affecting Pearl
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Moderator's Overview Biological Fac
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Introduction Downy mildew of pearl
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2 3 1 2 4 6 I 5 3 1 A s e x u a l s
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however, that wind plays an importa
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Figure 4. Figure assembly to demons
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in sterilized distilled water. Leav
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Frederiksen, R.A., and Rosenow, D .
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Tasugi, H. 1933. Studies on Japanes
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(Decarvalho 1949), Nigeria (King an
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Table 1. Differential and stable do
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ness against certain Phycomycetes (
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Large s c a l e f i e l d s c r e e
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References A I C M I P ( A U India
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Sun, M . H . , Chang, S.S., and Tsa
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Introduction Ergot (Claviceps fusif
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antidotale (Thakur and Kanwar 1978)
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Table 1. Performance of some select
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Table 4. Performance of some select
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Siddiqui, M . R . , and Khan, I . D
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184
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186
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188
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Sahelian Zone Tunisia 0 km 1000 Mor
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eflexa, and other scarab beetle spe
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200 Partially Burning, Bagging, and
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Table 1. Predators, parasites, and
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References Appert, J. 1957. Les Par
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Role and Utilization of Microbial A
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located through the root phloem. In
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They are not strong competitors in
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Fred, E.B., Baldwin, I . L . , and
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Climatic and Edaphic Limitations to
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much larger, about 2-4 kPa. Differe
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over the range that occurs in the f
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224 top 2 m holds from 100-250 mm o
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The conservative nature of qD is ex
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228 Table 4. Root and shoot charact
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Research on all these responses sho
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Making Millet Improvement Objective
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Table 1. Percentage of cultivated a
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• Well adapted, early-maturity, l
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in tests conducted by ICRISAT on fa
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ing a relatively good year in the S
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stress, as well as to genetic diffe
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Norman, D . W . , Newman, M . D . ,
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919.0 1212.0, Total area: 11.19 m i
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Table 4. Economics of pearl millet
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Azospirillum as a Seed Inoculant Az
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Management Practices to Increase Yi
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1500 1300 1100 900 700 500 300 100
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a physical form similar to SSP and
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Tillage The beneficial effects of t
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tivars of different maturity groups
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as fertility. The nitrogen contribu
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Greenland, D.J. 1958. Nitrate fluct
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Breeding for Adaptation to Environm
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15 S i k a r D i s t r i c t 15 Nia
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Table 2. Percentage of variation in
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Table 3. Correlation of the drought
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Selection for Traits Correlated to
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Discussion Squire's paper drew on d
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Androgenesis and Enzymatic Diversit
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Research on Pearl Millet Hybrids in
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La couleur des grains est variable.
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processed in different ways dependi
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Genetic Divergence in Landraces of
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Pearl Millet Regional Trials by CIL
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cultural practices such as early pl
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Host-Plant Resistance to Insect Pes
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of synthetics and composite varieti
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Pearl Millet Microbiology Potential
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available phosphate level in the so
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Pearl Millet Production in Drier Ar
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with cowpea system was the most eff
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Pearl Millet Pathology Disease Inci
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diseases hitherto undescribed are c
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Stunt and Counter-Stunt Symptoms in
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promising sources of resistance: Se
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69-188 mm for total seedling length
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un certain nombre de contraintes d'
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Priorities for Crop Protection Rese
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• those falling into other discip
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Striga Breeding for resistance to S
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ting improved cultivars to existing
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Survey of Pearl Millet Production a
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Table 4. Major constraints to produ
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Table 6. Extent of cultivation of r
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Table 10. Area planted to released
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Table 12. Production area and produ
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Participants Botswana Louis Mazhani
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Madhya Pradesh G.S. Chauhan Millet
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M . N . Prasad Professor & Head Cot
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S.K. Manzo Plant Pathologist Depart
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ICRISAT Center S. Appa Rao Botanist
Page 367 and 368:
RA-00110
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