PART II: Sorghum Genetic Enhancement Process

Conservation, Utilization and 3Distribution of Sorghum GermplasmN Kameswara Rao, PJ Bramel, V Gopal Reddy and UK Deb

Conservation, Utilization and 3Distribution of Sorghum GermplasmN Kameswara Rao, PJ Bramel, V Gopal Reddy and UK Deb

43

Conservation, Utilization and Distributionof Sorghum Germplasm

N Kameswara Rao1, PJ Bramel1, V Gopal Reddy2 and UK Deb3

3.1. IntroductionCollection, characterization and maintenance of plant genetic resources are essential for cropimprovement. ICRISAT has given high priority to this. The need to collect and conserve geneticdiversity in sorghum was realized some 35 years ago as traditional landraces became vulnerable toloss due to several factors: increasing adoption of modern varieties and hybrids; substitution ofsorghum by more remunerative crops, especially with extension of irrigation facilities as in Indiaand frequent and prolonged droughts in Africa. Thus, several landraces once abundant in parts ofAfrica and Asia are reported to be extinct now. The landraces Hegari, Zera-zera and Kurgis oncepresent in the Gezira, Kasala and Blue Nile provinces of Sudan, are on the verge of extinction or areno longer cultivated (Prasada Rao and Mengesha 1981a).

3.2. Germplasm Collection and AssemblyThe first major attempt toward assembling a world collection of sorghum was made by the IndianAgricultural Program of the Rockefeller Foundation (Murty et al. 1967; Rockefeller Foundation1970), and 16, 138 accessions were assembled from the major sorghum growing areas. Thiscollection was acquired by ICRISAT in 1974 through the All India Coordinated SorghumImprovement Project (AICSIP), Rajendranagar (Mengesha and Prasada Rao 1982). However, only8961 of the accessions were viable; the remainder had lost viability due to lack of proper storagefacilities before they were transferred. ICRISAT later obtained some 3000 of the missingaccessions from the duplicate sets maintained in the USA (Purdue and Fort Collins) and PuertoRico (Mayaguez) (Mengesha and Prasada Rao 1982). In 1974, in accordance with therecommendations of the Advisory Committee on Sorghum and Millets Germplasm sponsored bythe International Board for Plant Genetic Resources (IBPGR, now International Plant GeneticResources Institute, IPGRI), ICRISAT assumed the responsibility to maintain sorghum germplasmand to enlarge the world collection. Special efforts were made to collect or assemble landraces andwild relatives from areas threatened by genetic erosion. Between 1975 and 1997, ICRISATlaunched 32 collection missions in major sorghum growing areas and collected 8898 accessions(Table 3.1). Apart from ICRISAT’s own efforts, donations from several international and nationalorganizations such as IPGRI, Institut français de recherche scientifique pour le développement encoopération (ORSTOM), the Ethiopian Sorghum Improvement Program, Gezira Agricultural

1. Formerly of the International Crops Research Institute for the Semi-Arid Tropics, Patancheru 502 324, Andhra Pradesh, India.2. International Crops Research Institute for the Semi-Arid Tropics, Patancheru 502 324, Andhra Pradesh, India.3. Centre for Policy Dialogue, House 40/C, Road no. 11, Dhanmondi, Dhaka 1209, Bangladesh.

Kameswara Rao N, Bramel PJ, Gopal Reddy V and Deb UK. 2004. Conservation, utilization and distribution of sorghum germplasm.Pages 43-62 in Sorghum genetic enhancement: research process, dissemination and impacts (Bantilan MCS, Deb UK, Gowda CLL,Reddy BVS, Obilana AB and Evenson RE, eds.). Patancheru 502 324, Andhra Pradesh, India: International Crops Research Institute forthe Semi-Arid Tropics.

44

Research Station (Sudan), AICSIP, several Indian agricultural universities and individualscontributed to the collection (Prasada Rao et al. 1989). With 36, 774 accessions conserved from 91countries (Table 3.2), the ICRISAT Gene Bank now serves as a major repository of sorghumgermplasm. More than half of this collection is from five countries: India, Ethiopia, Sudan,Cameroon and Yemen. Although diversity from many countries has been adequately sampled,some remain poorly represented and are therefore high priority for future collection efforts. Thesecountries include Angola, Chad, China, Eritrea, Ivory Coast, Mozambique, Thailand, Turkey andZaire. Ethiopia, Indonesia, Myanmar, the Philippines and parts of West Africa are high-prioritycountries for collection of wild relatives (Stenhouse et al. 1997).

Table 3.1. Sorghum germplasm collected from various countries by ICRISAT expeditions undertaken from1975 to 1996.Country 1975-79 1980-84 1985-89 1990-961 Total

Botswana - 133 29 - 162Burundi - 72 - - 72Cameroon - 96 547 - 643Central African Republic - - 210 - 210Ethiopia - 131 - - 131Gambia - 56 - - 56Ghana - 83 - - 83India 652 267 947 734 2600Kenya 2 - - - 2Lesotho - - 249 - 249Malawi 288 6 - - 294Mali - - - 20 20Mozambique - 48 - - 48Myanmar - 4 - - 4Namibia - - - 181 181Nigeria - 200 - 184 384Pakistan - - 46 - 46Russia (former) - - - 41 41Rwanda - 73 - - 73Sierra Leone - 104 - - 104Somalia 120 - 271 - 391South Africa - 63 - - 63Sri Lanka - 23 - - 23Sudan 130 30 - - 160Swaziland - - 182 - 182Tanzania 107 44 267 - 418Togo - - 34 - 34Uganda - - - 448 448Yemen - 163 - - 163Zambia - 204 - - 204Zimbabwe - 228 1131 - 1359Total 1299 2028 3913 1658 88981. There was no collection expedition in 1995.

45

Table 3.2. Source of the sorghum germplasm assembled at the ICRISAT Gene Bank, 2002.Country/institution Number of accessions Country/institution Number of accessionsAsia AfricaAfghanistan 5 Senegal 241Bangladesh 9 Sierra Leone 108China 646 Somalia 446ICRISAT 548 South Africa 937India 6202 Sudan 2505Indonesia 36 Swaziland 203Iran 17 Tanzania 718Iraq 3 Togo 294Israel 22 Uganda 1763Japan 108 Zaire 52Korea, Republic of 78 Zambia 362Lebanon 360 Zimbabwe 1576Maldives 10Myanmar 20 AmericasNepal 8 Argentina 18Pakistan 88 Colombia 4Philippines 61 Cuba 2Russia (former) 405 Dominican Republic 2Saudi Arabia 22 El Salvador 2Sri Lanka 25 Guatemala 7Syrian Arab Republic 4 Haiti 1Taiwan 6 Honduras 65Thailand 9 Jamaica 3Turkey 51 Mexico 83Yemen, Republic of 2164 Nicaragua 2

USA 2128Africa Uruguay 1Algeria 23 Venezuela 173Angola 44Benin 199 EuropeBotswana 219 Belgium 1Burkina Faso 548 Cyprus 1Burundi 140 France 5Cameroon 2485 Germany 7Cape Verde 1 Greece 1Central African Republic 249 Hungary 87Chad 193 Italy 9Congo 1 Portugal 6Egypt 35 Romania 7Ethiopia 4400 Spain 3Ghana 147 United Kingdom 3Gambia 57Ivory Coast 7 OceaniaKenya 990 Australia 63Lesotho 271 Papua New Guinea 1Madagascar 14Malawi 423 Unknown 170Mali 694 Total number of accessions 36, 774Mauritania 16 Total number of countries 91Morocco 27Mozambique 48Namibia 189Niger 413Nigeria 1683Rwanda 291

46

3.3. Conservation and MaintenanceSorghum seeds show orthodox storage behavior (ie, desiccation tolerant); hence, the germplasm isconserved ex situ as seeds in the gene bank under controlled conditions. Ex situ conservation is costefficient, and also makes it easier for scientists to access, study, distribute and use plant geneticresources. At ICRISAT, all the germplasm is stored as active collection under medium-term storageconditions (4oC and 20% RH). For each accession, about 300-400 g of seed is harvested at optimummaturity, usually 6-7 weeks after anthesis, from postrainy-season multiplication plots. The seedsare dried in the shade to about 8% moisture content, threshed by hand, cleaned and stored inscrew-capped aluminium containers. The germplasm is also conserved as base collection at -20°Cfor long-term storage. For this purpose, about 125 g of seed is cleaned and dried to 5-6% moisturecontent by equilibration with air at 20°C and 15% RH for approximately 3 weeks. The dried grainis vacuum-sealed in aluminum foil bags and stored after confirming that initial viability is more than95%. Seeds stored under medium-term conditions (active collection) are used for distribution andresearch, while those stored under long-term conditions (base collection) are used only forregenerating active collections. Viability is monitored by germination tests at 5-year intervals inmedium-term storage and 10-year intervals in long-term storage. Any sample having less than 75%viability is identified for rejuvenation.

The medium- and long-term storage rooms are constructed on a modular principle withfabricated panels. Each room has a capacity of 125 m3, capable of accommodating over 30,000germplasm accessions. The storage rooms have mobile shelving systems, and are equipped withstandby refrigeration and dehumidification systems. The temperature and relative humidity insidethe storage rooms are monitored regularly with thermohygrographs. Audible and visual electronicalarm systems have been installed to safeguard the seeds from any rise in temperature and relativehumidity and to help maintain the desired conditions. The Gene Bank has a power generator tocope with long periods of power failure.

Chapter 14G of Agenda 21 of the United Nations Conference on the Environment andDiversity (UNCED) recommends that all gene banks duplicate their collections of germplasm forsafety under long-term conditions. ICRISAT’s agreement with the Food and AgricultureOrganization of the United Nations (FAO) also requires safety duplication. ICRISAT has amemorandum of understanding for the duplication of sorghum germplasm with the Southern AfricanDevelopment Community (SADC), Regional Gene Bank (SRGB), Zambia, and recently transferred4527 accessions of African origin. Efforts are being made to identify sites for the safe duplication ofthe complete collection. Once the sites are identified, agreements will be made to facilitate transferand safe storage of the duplicate collection. Other options for duplication are also being explored. Forexample, the sorghum germplasm collections at ICRISAT and at the National Plant GermplasmSystem (NPGS), USA, overlap considerably and cross-referencing the two databases revealed thatover 21,985 accessions (approximately 60% of ICRISAT holdings) are duplicated. Obtaining uniqueaccessions from NPGS to add to the world collection at ICRISAT and duplication of those notavailable at NPGS are being considered.

Sorghum is a partially outbreeding and self-pollinated species. Therefore, germplasmaccessions are maintained by selfing. During collection and initial multiplication at ICRISAT,variant types were separated so that each accession is a true-breeding single type. To preventcontamination by foreign pollen, individual panicles are covered with selfing bags as soon as theyemerge from the boot prior to anthesis. They are kept covered for at least 3 weeks, after which the

47

bags are removed to assist in seed drying and maturity. Seed from at least 30 selfed plants is bulkedto maintain an accession. Wild relatives are also maintained by selfing. However, a number of speciesdo not set seeds and are maintained as live plants in a field gene bank (Stenhouse et al. 1997).

Diversification, genetic studies and germplasm enhancement through conversion andintrogression resulted in the development of trait-specific germplasm pools over the years atICRISAT. These, along with the germplasm accessions possessing some valuable traits, aremaintained as subsets or working collections for ready access and use. In addition to the activecollections, the following working collections were established and maintained at ICRISAT on therecommendation of various sorghum workers (Prasada Rao et al. 1989).

Spontaneous collection. This includes 542 wild and weed races from 33 taxa (Table 3.3).

Named cultivar collection. This includes 237 named cultivars released by private and publicinstitutions in different countries. About 2 kg seed samples are maintained to meet seed requests.

Table 3.3. Wild relatives of sorghum assembled at ICRISAT until December 1999.Genus/section/species/subspecies Race Number of accessions

Genus: Sorghum arundinaceum 33Section: Sorghum aethiopicum 16S. bicolor/arundinaceum lanceolatum 7

macrochaeta 3rhizomatous 2usumbaranse 3verticilliflorum 87virgatum 18

S. bicolor/drummondii drummondii 153hewisonii 2

S. halepense halepense 22controversum 4miliaceum 5almum 6

S. propinquum 3Section: Chaetosorghum S.macrospermum 1Section: Heterosorghum S. laxiflorum 1Section: Parasorghum S. australiense 3S. brevicallosum 1S. matarankense 2S. purpureosericeum 2

deccanense 4dimidiatum 2

S. versicolor 6Section: Stiposorghum S. affstipodeium 2S. Intrans 5S. plumosum 1S. stipoideum 5Section: Unclassified Para sorghum 10Others 2Genus: Sorghastrum 6Total 417

48

Genetic stock collection. This consists of genotypes resistant to diseases and pests, lines withidentified genes and cytoplasmic-genetic male steriles. Seed samples of 1 kg are maintained byselfing, except in the case of male-sterile lines, which are maintained by hand pollination.

Conversion collection. To augment the use of tropical sorghum germplasm in breeding programs,and to broaden the genetic base, ICRISAT began a program to convert the tall, photoperiod-sensitive landraces into photoperiod-insensitive lines. Currently, the collection has 176 conversionlines obtained from USA and 348 lines developed at ICRISAT, mostly from the Zera-zera landraces(from Ethiopia and Sudan), which are highly prized for their superior agronomic characteristics butare of restricted utility because of their photoperiod sensitivity and plant height.

Core collection. A sorghum core collection consisting of 3,475 accessions was established afterstratifying the total world collection geographically and taxonomically into subgroups followed byclustering into closely related groups on the basis of characterization data using principle componentanalysis. Representative accessions from each cluster were drawn in proportion to the total number ofaccessions in the subgroup (Prasada Rao and Ramanatha Rao 1995). This core collection representsthe genetic spectrum of the whole collection for utilization in crop improvement.

3.4. Characterization and EvaluationCharacterization of germplasm facilitates its utilization for crop improvement. Much of the germplasmassembled at ICRISAT was characterized for important morpho-agronomic characters at Patancheruduring both the rainy and postrainy seasons, using the crop descriptors co-published by IBPGR andICRISAT (IBPGR/ICRISAT 1993). The range of variation observed in the collection for 21 importanttraits is summarized in Table 3.4. Substantial variation could be seen in days to flowering (very early 33days to very late 199 days), plant height (less than 1 m to more than 6 m), panicle size (head length 2.5-90 cm and head width 1.0-80 cm), shape, plant pigmentation, grain color and grain mass.

Traditional landraces and wild relatives evolve through centuries of introgression, andnatural and human selection and acquire resistance to specific pests, diseases and environmentalstresses, which can be used as sources of resistance. Over the years, ICRISAT scientists havescreened sorghum germplasm accessions for resistance to diseases and pests under natural as wellas artificial conditions (Stenhouse et al. 1997). For example, screening of germplasm accessionsresulted in the identification of 60 accessions resistant to shoot fly, 169 accessions tolerant tostem borer and 12 accessions resistant to midge. Stable resistance to both shoot fly and stemborer was found in IS 1082, IS 1855, IS 2122, IS 2134, IS 2205, IS 2312, IS 5470, IS 5480, IS5604, IS 18554 and IS 18577. Similarly, sources of resistance to grain molds were identified inmore than 156 accessions, resistance to anthracnose was found in 14 germplasm accessions andresistance to downy mildew in 89 accessions. Germplasm accessions that have resistance to morethan one disease were also identified, eg, IS 18758 (anthracnose and rust); IS 3443, IS 18882(downy mildew and rust); IS 17141 (grain mold and anthracnose) and IS 3547, IS 8283 (grainmold, downy mildew and rust).

The germplasm has also been screened for resistance to Striga, abiotic stresses (drought andtemperature stress) and for capability of emergence at higher soil temperatures. Some germplasmaccessions identified as resistant to Striga and used in breeding programs are IS 2221, IS 4202, IS5106, IS 7471, IS 8741, IS 9830, IS 9951 and IS 18331. The results of screening conducted overthe years at ICRISAT and the number of promising lines identified with 20 useful traits aresummarized in Table 3.5.

49

The morphological variation in some accessions was found to be associated with economicallyimportant traits. For example, the presence of a phenolic compound (flavan-4-ol) in the pericarpresults in a red pigmentation that is associated with resistance to grain molds. Of the 7123accessions screened for grain mold resistance, all but one of the 156 resistant lines identified havea colored pericarp (Bandyopadhyay et al. 1988). Similarly, tan plant color is associated withresistance to leaf diseases and grain weathering. A brown midrib appears to be associated with lowlignin content and high digestibility of plant parts. The dull green midrib color is used to identifysweet-stalk sorghums for chewing in South Africa (Stenhouse et al. 1997).

In addition to resistance to biotic and abiotic stresses, other important genes have also beenobtained from the germplasm. For example, Singh and Axtel (1973) reported a high-lysine gene fromtwo Ethiopian sorghums (IS 11167 and IS 11758), associated with relatively high levels of protein.Prasada Rao and Murthy (1979) reported the collection of basmati (scented) sorghums in India.

Multilocational evaluation of germplasm by ICRISAT scientists in collaboration withscientists of national programs in India, Kenya, Nigeria, Somalia and Thailand led to theidentification of locally-adapted varieties and thus broadened the sorghum genetic base in thesecountries. Table 3.6 shows the list of collaborators, locations, number of accessions and the type ofgermplasm evaluated during different years. For instance, in a collaborative effort with theNational Bureau of Plant Genetic Resources/Indian Council of Agricultural Research (NBPGR/ICAR), India, accessions originating from 35 countries, but predominantly from Ethiopia, India andUganda, were evaluated for 26 characters at 3 locations to identify useful and adapted lines. Theresults were published as a catalog (Mathur et al. 1992).

Table 3.4. Range of variation in selected characteristics of sorghum germplasm.Characteristics Range of variation

Plant height (postrainy season) 50-580 cmPlant height (rainy season) 65-655 cmPlant pigmentation Tan to pigmentedNumber of basal tillers 1-20Nodal tillering Present to absentMidrib color White to brownDays to flowering (postrainy season) 36-154Days to flowering (rainy season) 33-199Panicle exertion 0-72 cmPanicle length 2.5-90 cmPanicle width 1-80 cmPanicle compactness and shape Very loose, stiff branches to compact ovalGlume color Straw to blackGlume covering Fully covered to uncoveredSeed color Straw to blackSeed lustre Lustrous to nonlustrousSeed subcoat Present to absentSeed size 0.8-8 mm100-seed weight 0.29-8.92 gEndosperm texture Completely starchy to completely corneousThreshability Freely threshable to difficult to thresh

50

Tabl

e 3. 5

. Sou

rce o

f use

ful t

raits

1 iden

tified

from

the s

orgh

um g

erm

plas

m m

ainta

ined

at IC

RISA

T Ce

nter

.Tr

aits

Char

acter

istics

Coun

tryA

BC

DE

FG

HI

JK

LM

NO

PQ

RS

TTo

tal

Alge

ria1

1An

gola

11

2Ar

genti

na4

4Au

strali

a9

615

Benin

11

Botsw

ana

220

11

24Bu

rkina

Fas

o1

61

15

14Ca

mero

on10

13

18

104

766

111

1Ce

ntral

Afric

an R

epub

lic1

1Ch

ad3

24

9Ch

ina15

51

115

37Do

minic

an R

epub

lic1

1Eg

ypt

45

110

Ethio

pia16

103

18

13

54

23

56Fr

ance

11

Germ

any

11

Ghan

a11

314

533

Hung

ary

11

India

195

348

200

4142

16

11

1111

426

365

137

5810

1715

46Ind

ones

ia1

1Ira

n1

12

Israe

l1

89

Italy

11

Jama

ica2

2Ja

pan

61

31

11Ke

nya

17

31

51

101

21

32Ko

rea,

Repu

blic o

f1

1Le

bano

n1

5415

13

74Le

sotho

5656

Malaw

i5

41

212

Mali

81

110

Mexic

o8

11

313

...co

ntin

ued

51

Tabl

e 3. 5

. Con

tinu

ed

Trait

sCh

arac

terist

ics

Coun

tryA

BC

DE

FG

HI

JK

LM

NO

PQ

RS

TTo

tal

Myan

mar

11

2Ne

pal

11

Nige

r3

25

Nige

ria17

110

13

11

151

341

410

57

111

Pakis

tan1

61

8Po

rtuga

l2

2Se

nega

l1

21

15

Soma

lia6

17

South

Afric

a15

81

242

12

319

1Sr

i Lan

ka1

1Su

dan

131

68

55

219

37

102

1337

43

1311

459

Swaz

iland

272

29Sy

rian A

rab R

epub

lic1

1Ta

iwan

11

Tanz

ania

21

12

915

Thail

and

21

3Tu

rkey

51

1622

USSR

(for

mer)

41

16

Ugan

da5

82

21.0

04

23

231

33

820

85Un

ited K

ingdo

m1

1US

A7

305

35

325

511

122

202

96

2362

500

Yeme

n, Re

publi

c of

81

7180

Zaire

11

Zamb

ia1

59

15Zim

babw

e90

17

21

310

4Un

know

n6

12

11

112

413

6To

tal23

634

814

1360

6912

615

614

3189

2051

236

7616

211

720

151

274

3883

1. A

= na

med

cultiv

ars;

B =

ICRI

SAT

conv

ersio

n lin

es; C

= ra

iny-se

ason

bas

ic co

llecti

on; D

= sh

oot f

ly re

sistan

t; E

= ste

m bo

rer;

F= m

idge;

G =

head

bug

; H =

gra

in mo

ld;I =

anthr

acno

se; J

= ru

st; K

= do

wny m

ildew

; L =

Strig

a res

istan

t; M =

gloss

y; N

= pop

sorg

hum;

O =

swee

t stal

k; P

= twi

n see

ded;

Q = l

arge

glum

e; R

= bloo

mles

s; S

= bro

omco

rnan

d T =

cytop

lasmi

c male

-ster

ile.

52

Table 3.6. Collaborators1, locations, number of accessions and type of sorghum germplasm materialcharacterized/evaluated by ICRISAT in different years.Year Collaborator Location Number of accessions Type of material

1986 NBPGR, India Hisar 2000 Photoperiod sensitiveNBPGR, India Trichur 2000 Photoperiod sensitiveNBPGR, India Issapur 1500 Forage sorghumsNBPGR, India Hisar 1500 Forage sorghumsNBPGR, India Jhansi 1500 Forage sorghumsNBPGR, India Akola 1500 Forage sorghums

1987 NBPGR, India Trichur 2000 Photoperiod sensitiveNBPGR, India Akola 99 Evaluation for midseason drought resistanceNBPGR, India Issapur 4577 Entire Indian collection and promising forage types

1988 NBPGR, India Issapur 2993 Selected germplasmAICSIP, India Akola 1305 Rainy-season basic collectionAICSIP, India Coimbatore 1305 Rainy-season basic collectionAICSIP, India Indore 1305 Rainy-season basic collectionAICSIP, India Rajendranagar 1305 Rainy-season basic collectionAICSIP, India Surat 1305 Rainy-season basic collectionMinistry of Bonka, Baidoa 2066 Rainy-season short-duration sorghumsAgriculture, Somalia

1989 NBPGR, India Issapur 1200 Selected germplasm for forageIGFRI, India Jhansi 1200 Selected germplasm for forageNRCS, India Rajendranagar 1200 Selected germplasm for forageNBPGR, India Akola 1000 Selected germplasm for grainTNAU, India Coimbatore 1000 Selected germplasm for grainNRCS, India Rajendranagar 1000 Selected germplasm for grainUAS, India Bijapur 1002 Selected germplasm for postrainy-season programMPKV, India Mohol 1002 Selected germplasm for postrainy-season programNRCS, India Rajendranagar 1002 Selected germplasm for postrainy-season programWASIP, Nigeria Bagauda 2010 Early-maturing germplasm

1990 NBPGR, India Issapur 500 Selected germplasm for forageIGFRI, India Jhansi 500 Selected germplasm for forageNRCS, India Rajendranagar 500 Selected germplasm for forageNBPGR, India Akola 500 Selected germplasm for grainTNAU, India Coimbatore 500 Selected germplasm for grainNRCS, India Rajendranagar 500 Selected germplasm for grainMPKV, India Rahuri 200 Selected germplasm for postrainy-season programUAS, India Bijapur 200 Selected germplasm for postrainy-season programICRISAT, India Patancheru 200 Selected germplasm for postrainy-season programWASIP, Nigeria Bougouda 70 Selected early-maturing germplasm for grainEARCAL, Kenya Kiboko 69 Selected germplasm for grain

1991 NBPGR, India Issapur 200 Forage sorghumsIGFRI, India Jhansi 200 Forage sorghumsNRCS, India Rajendranagar 200 Forage sorghumsNBPGR, India Akola 200 Forage sorghumsAICSIP, India Surat 200 Forage sorghumsNRCS, India Rajendranagar 200 Forage sorghumsNBPGR, India Patancheru 200 Forage sorghumsNRCS, India Rahuri 1200 Selected germplasm for postrainy-season programNRCS, India Sholapur 1200 Selected germplasm for postrainy-season program

...continued

53

3.5. DistributionICRISAT’s gene bank has become a major source of diversity available to sorghum plant breeders.So far, ICRISAT has distributed sorghum germplasm samples to users in 105 countries (Tables 3.7and 3.8). About 92% of the germplasm was distributed to national and international programs anduniversities, while 8% was provided to the private sector. The year-wise distribution of sorghumgermplasm (246,901 samples) to different regions is given in Table 3.8.

Table 3.6. Continued

Year Collaborator Location Number of accessions Type of material

NRCS, India Bijapur 1200 Selected germplasm for postrainy-season programNRCS, India Rajendranagar 1200 Selected germplasm for postrainy-season programNRCS, India Patancheru 200 Selected germplasm for postrainy-season programINTSORMIL/Purdue Wad Medani (Sudan) 2340 Sorghum germplasm of Sudanese originUniversity

1992 NBPGR, India Issapur 200 Forage sorghumsIGFRI, India Jhansi 200 Forage sorghumsNRCS, India Rajendranagar 200 Forage sorghumsICRISAT, India Patancheru 200 Forage sorghumsNBPGR, India Akola 200 Forage sorghumsAICSIP, India Surat 200 Forage sorghumsAICSIP, India Udaipur 200 Forage sorghumsNRCS, India Rajendranagar 200 Forage sorghumsICRISAT, India Patancheru 200 Forage sorghumsNRCS, India Rajendranagar 1200 Selected germplasm for rainy-season programNRCS, India Sholapur 1200 Selected germplasm for rainy-season programNRCS, India Bijapur 1200 Selected germplasm for rainy-season programNRCS, India Rajendranagar 1200 Selected germplasm for rainy-season program

1993 NBPGR, India Issapur 200 Forage sorghumsGAU, India Anand 200 Forage sorghumsNRCS, India Rajendranagar 200 Forage sorghumsICRISAT, India Patancheru 200 Forage sorghumsNBPGR, India Akola 200 Forage sorghumsAICSIP, India Surat 200 Forage sorghumsAICSIP, India Udaipur 200 Forage sorghumsRAU, India Durgapura 200 Forage sorghumsICRISAT, India Patancheru 200 Forage sorghumsFCRI, Thailand Suphan Buri 500 Forage, grain, dual purpose

1994 NBPGR, India Issapur 200 Forage sorghumsICRISAT, India Patancheru 200 Forage sorghumsNBPGR, India Akola 200 Forage sorghumsAICSIP, India Surat 200 Forage sorghumsAICSIP, India Indore 200 Forage sorghumsNRCS, India Rajendranagar 200 Forage sorghumsTNAU, India Coimbatore 200 Forage sorghumsICRISAT, India Patancheru 200 Forage sorghums

1. NBPGR = National Bureau of Plant Genetic Resources; AICSIP = All India Coordinated Sorghum Improvement Project; IGFRI = Indian Grassland and FodderResearch Institute; NRCS = National Research Centre for Sorghum; TNAU = Tamil Nadu Agricultural University; UAS = University of Agricultural Sciences;MPKV = Mahatma Phule Krishi Vidyapeeth, ICRISAT = International Crops Research Institute for the Semi-Arid Tropics; WASIP = West African SorghumImprovement Program; EARCAL = Eastern Africa Regional Cereals and Legumes Network; INTSORMIL = International Sorghum/Millet CollaborativeResearch Support Program; GAU = Gujarat Agricultural University; RAU = Rajasthan Agricultural University and FCRI = Field Crops Research Institute.

54

Table 3.7. Distribution of sorghum lines from ICRISAT’s Gene Bank.

Country 73-74 75-79 80-84 85-89 90-94 95-99 2000 2001 2002 Total

Argentina 0 83 213 110 5 0 411Australia 3 86 425 132 25 5 2 3 9 690Austria 0 0 98 0 0 0 98Burundi 0 48 25 217 0 0 290Belgium 0 22 0 1251 224 56 1553Benin 0 7 0 17 0 0 24Burkina Faso 0 359 2219 3640 0 0 6218Bangladesh 0 414 143 42 0 34 633Bolivia 0 24 0 50 0 0 74Brazil 352 131 159 1864 79 0 9 2594Barbados 0 0 91 0 0 0 91Botswana 0 75 1240 472 0 0 70 1857Cambodia 0 0 0 0 0 5 5Central African Republic 0 0 0 0 3 0 3Canada 2 52 0 14 350 106 524China 0 70 108 602 33 32 10 855Cameroon 0 0 2777 85 16 0 2878Chad 0 0 0 141 0 0 141Colombia 0 10 0 573 0 0 583Cape Verde 0 0 125 0 0 0 125Costa Rica 0 0 0 49 0 0 49Czechoslovakia 0 0 30 0 0 0 30Djibouti 0 0 0 50 0 0 50Denmark 0 0 10 0 0 0 3 13Dominican Republic 0 0 2 0 0 0 2Ecuador 0 0 0 42 8 0 50Egypt 0 782 0 147 1075 239 2243El Salvador 0 36 49 0 0 0 85Ethiopia 0 1751 651 656 31 0 452 3541Fiji 0 0 0 0 20 0 1 21France 0 33 649 342 189 290 7 6 10 1526Ghana 0 19 182 5 0 92 2 300Gambia 0 0 0 55 0 0 55Germany 0 4 2 419 38 1 6 470Guatemala 0 0 608 285 0 0 893Guyana 0 0 0 99 0 0 99Honduras 0 0 0 88 0 0 36 124Haiti 0 0 0 0 0 0 28 28Hungary 0 78 0 15 0 0 93Indonesia 0 0 2 0 78 98 178India 3 12145 10047 59352 27915 11648 638 2265 1626 125639Ireland 0 12 0 0 0 0 12Iran 0 0 0 620 425 12 16 1073Iraq 0 0 0 28 0 0 28Israel 0 0 20 100 0 0 120Italy 0 55 166 777 239 10 10 1257Jamaica 0 0 0 0 426 0 426Jordan 0 0 0 0 0 5 5

...continued

55

Table 3.7. Continued

Country 73-74 75-79 80-84 85-89 90-94 95-99 2000 2001 2002 Total

Japan 0 0 668 244 73 9 3 997Kenya 0 1132 2426 1471 5945 68 11042Korea, Republic of 0 170 0 932 0 0 81 1183Kuwait 0 0 0 0 0 0 2 2Lebanon 0 105 0 0 0 0 105Liberia 0 0 0 50 0 0 50Libyan Arab Jamahiriya 0 20 0 0 70 0 90Morocco 0 0 231 0 0 0 231Mexico 0 1040 329 3244 105 0 4718Mali 0 189 1279 424 559 1273 314 4038Mozambique 0 0 0 248 0 0 6 254Mauritania 0 0 33 0 0 0 33Malawi 0 57 1244 509 0 0 1810Myanmar 0 0 0 8 0 0 8Malaysia 0 101 0 0 0 0 101Niger 0 28 369 3 1838 15 1 2254Nigeria 0 366 101 2075 415 20 2977Nicaragua 0 0 0 46 0 0 46Netherlands 0 41 99 6 2 0 148Nepal 0 5 0 0 20 0 25New Zealand 0 0 2 0 0 0 2Pakistan 0 49 276 37 2 15 17 396Peru 0 45 0 0 16 0 61Philippines 0 7 131 207 26 0 371Papua New Guinea 0 49 83 6 0 0 138Qatar 0 126 30 202 0 0 358Romania 0 0 114 0 20 0 134Russia (former) 0 422 442 35 549 0 2 1450Rwanda 0 81 161 1052 99 0 1393Saudi Arabia 0 57 0 71 78 0 206Sudan 0 1184 1049 0 2794 181 5208Senegal 0 19 29 183 0 0 231Solomon Islands 0 0 0 0 0 0 25 25Sierra Leone 0 0 35 118 30 0 50 233Somalia 0 59 839 2159 0 0 3057South Africa 0 0 23 0 132 29 184Sri Lanka 4 36 21 2 0 35 98Surinam 0 37 8 0 0 0 45Switzerland 0 0 0 67 1 0 68Syrian Arab Republic 0 0 98 0 21 2 121Thailand 0 0 158 10 820 214 1202Trinidad and Tobago 0 0 0 0 0 15 15Turkey 0 0 0 99 0 50 149Taiwan 0 25 41 17 0 0 83Tanzania 0 329 1290 0 0 0 40 1659United Arab Emirates 0 0 0 0 0 0 168 87 255United Kingdom 1 123 702 47 155 36 3 1067Uganda 0 0 903 46 1369 0 2318

...continued

56

Table 3.8. Year-wise distribution of sorghum germplasm to different regions (as on December 2002).Year Africa America Asia Europe Oceania Total

1973 3 31974 354 7 1 3621975 148 750 2255 15 31681976 1682 146 2363 89 42801977 2097 503 3126 61 135 59221978 473 72 2164 27 27361979 2175 343 3958 176 66521980 268 461 2129 207 390 34551981 2616 1343 3755 38 16 77681982 3326 4839 2483 128 10,7761983 9476 284 2922 802 21 13,5051984 5444 5899 1133 747 83 13,3061985 7474 573 3731 1382 18 13,1781986 2715 3780 15427 1075 20 23,0171987 4489 519 13456 300 18,7641988 5625 1406 20630 29 99 27,7891989 842 3250 11369 237 1 15,6991990 4051 325 7419 133 40 11,9681991 2553 47 7124 397 10,1211992 878 4067 5475 222 5 10,6471993 6772 7 7468 26 14,2731994 3652 875 2736 90 73531995 682 187 1992 20 5 28861996 92 34 2903 15 30441997 822 3 3023 52 39001998 110 86 3486 340 40221999 245 729 170 11442000 365 76 1054 26 2 15232001 495 2355 1005 3 38572002 80 17 1639 22 5 1783

Total 69,646 30,246 138,311 7832 866 246,901

Table 3.7. Continued

Country 73-74 75-79 80-84 85-89 90-94 95-99 2000 2001 2002 Total

Uruguay 0 0 126 0 30 0 156USA 0 316 11228 3041 4046 178 7 990 17 19823Venezuela 0 40 13 23 256 11 5 348Vietnam 0 0 0 0 4 0 4Yemen, Republic of 0 134 237 2104 178 29 132 2814Yugoslavia (former) 0 0 52 0 0 154 206Zaire 0 10 0 2149 0 0 2159Zambia 0 60 1240 383 3431 0 5114Zimbabwe 0 0 2659 4790 99 29 4 7581Total 365 22758 48810 98447 54362 14996 1523 3857 1783 246,901

57

ICRISAT has responded positively to requests from national programs for the restoration oflost or damaged germplasm. The germplasm lost during the civil wars in Ethiopia and Rwanda wasrestored from the sorghum collections maintained in ICRISAT’s Gene Bank. Similarly, sorghumcollections from Botswana, Cameroon, Kenya, Nigeria and Somalia were repatriated to thosecountries for conservation and utilization (ICRISAT 1991).

As part of the FAO International Network of Ex Situ Collections, ICRISAT designated over80% of the sorghum collection to the auspices of FAO/CGIAR. The agreement covers collectionsheld by ICRISAT prior to December 1993, when the Convention on Biological Diversity (CBD)which affirmed sovereign rights of national governments over their natural resources came intoeffect. Though the designated germplasm will continue to be readily available to all, ICRISAT willhave to ensure that recipients do not apply for intellectual property rights to it. For this, ICRISATrequires every recipient of germplasm to sign a Material Transfer Agreement (MTA). Theavailability of germplasm acquired after December 1993 is subject to conditions imposed by thesource country.

3.6. DocumentationThe morphological and agronomic data along with the passport information were until recentlydocumented and stored on a VAX-based computer program called ICRISAT Data Managementand Retrieval System (IDMRS) written in BASIC language. However, since information retrievalwas slow, it was first replaced with System 1032 of ‘CompuServe Technology’ and now withMicrosoft Access, a Relational Database Management System (RDBMS). MS Access facilitates amore efficient management of large databases and quicker retrieval of information in full or in partand in combinations of desirable traits to suit specific requirements. ICRISAT has supplied manycopies of the passport and characterization database to scientists in India, Cameroon, Chad, China,Ethiopia, Germany, Mexico, UK and the USA (Prasada Rao et al. 1989).

Access to information on collections facilitates greater utilization of genetic diversity.ICRISAT is involved in the System-wide Information Network for Genetic Resources (SINGER)by which information on collections of genetic resources maintained at CGIAR centers can beaccessed and searched collectively. SINGER provides ready access on the identity, origin andcharacteristics of the accessions in ICRISAT’s Gene Bank to users worldwide through a website onthe Internet.

3.7. UtilizationAs with most crops, the earliest phase of sorghum improvement in Africa and Asia utilized pure lineselections from among and within indigenous landraces, exploiting variations that arose frommutations and natural hybridization between different plant types. The second phase involveddeliberate hybridization between landraces (Stenhouse et al. 1997). Some of the importantderivatives of landraces include Maldandi in India, which still occupies millions of hectares in thepostrainy-season sorghum belt; Naga White (IS 17632) in Ghana, notable for its exceptional seedlingvigor; Segaolane (IS 18535) in Botswana; Framida (IS 8744), selected in South Africa from a varietyfrom Chad and notable for its excellent resistance to Striga asiatica; Serena (IS 18520) in Tanzaniaand Uganda, subsequently shown to have some resistance to Striga and shoot fly, and found to beresistant to bird damage because of its high-tannin brown grain (Stenhouse et al. 1997).

58

The Zera-zera landraces from Ethiopia and Sudan have proved to be useful sources of manytraits such as excellent grain quality, high grain yield potential, tan plant, straw glume color,resistance to leaf diseases, tolerance to grain weathering and desirable plant type (Prasada Raoand Mengesha 1979, 1981b). So extensive was their use in sorghum breeding programs atICRISAT and national sorghum breeding programs, that they dominate the germplasm base ofthe elite materials. Other groups of germplasm have also been used as sources of specific traits.Indian landraces (Durra) have provided sources of resistance to drought, shoot fly, stem borerand midge. The Kafir race of sorghum in combination with Durra sorghums from eastern Africaprovided the basis of the nuclear cytoplasmic male-sterility system that is the basis of allcommercial hybrid sorghum breeding. The Guinea race from West Africa has providedresistance to grain mold and Bicolor sorghums, which have contributed to the breeding of foragesorghums. In some cases, germplasm accessions were directly released as improved cultivars. Forexample, E 35-1 (a selection from a Zera-zera landrace from Ethiopia) was recommended forrelease in Burkina Faso , and IS 9302 and IS 9323 (Kafir landraces from South Africa) werereleased for intermediate altitude areas of Ethiopia (Prasada Rao et al. 1989). Table 3.9 lists thenumber of sorghum germplasm accessions or selections released to date as superior varieties indifferent countries.

Alternative uses of sorghum. Normally, sorghum grain is ground or pounded after removing thepigmented pericarp. The flour is used to make porridge, bread or beer. Alternative uses of sorghumare becoming increasingly important and its use as food is slowly declining. For example, there is anincreasing demand for forage-sorghum genotypes. Breeders of such genotypes in India havescreened the world collection and identified many promising plant types with desirable attributeslike plant height, profuse leafiness and dry matter digestibility. In a joint evaluation with NBPGR inIndia, 165 promising germplasm lines were identified for forage improvement (Mathur et al. 1991,1992).

Sorghum grain and stalk too have vast potential for industrial utilization. Technology is nowavailable to produce sugar, alcohol, starch, semolina and malt products from sorghum grain. Large-scale urbanization in Africa has resulted in the transformation of sorghum beer production from acommunity affair to an industrialized process (opaque beer brewing). Most reddish-brownsorghums from the Greater Horn of Africa and Southern Africa (Rwanda, Zimbabwe, Burundi andEthiopia) are used for making beer. In West Africa, 75-90% of the sorghum grown (white grained)is used to make thin and thick porridges. In China, sorghum is mainly used to prepare a popularalcoholic drink (sorghum wine), vinegar and as poultry feed.

Sorghum grain is a major component of cattle, pig and chicken feeds in the USA, Central andSouth America, Australia and China. The grain is usually processed by grinding, rolling, flaking orsteaming to improve digestibility. Dried sorghum stems that remain standing in the field afterharvest are often grazed or cut and fed to livestock (Stenhouse et al. 1997). The fibrous stem is alsoused to manufacture fibre boards for construction purposes.

Sorghum landraces with sweet stalks are sparingly distributed across sorghum growing areasin Africa and India. The green and tender stalks are chewed like sugarcane. In Ethiopia, sweet-stalksorghums are also used to make confectionery and to produce alcohol, adopting the sugarcanetechnology. Screening part of the world collection of sorghum germplasm for stalk sugar contentrevealed that 78 promising lines had sugar content ranging from 16.2-38.1% (on dry weight basis)(Subramaniam et al. 1987).

59

Table 3.9. Sorghum germplasm accessions or selections released as superior varieties in differentcountries.Accession number Country of origin Country of release Released name Year of release Remarks

IS 8965 Kenya Myanmar Shwe-ni 1 1980IS 2940 USA Myanmar Shwe-ni 2 1981IS 18758 Ethiopia Burkina Faso E 35-1 1983IS 23520 Ethiopia Zambia Sima 1989IS18758 Ethiopia Burundi Gambella 1107 1990IS 18484 India Honduras Tortillerio 1 1984IS 302 China Myanmar Shwe-ni 10 1980IS 5424 India Myanmar Shwe-ni 8 1980IS 8571 Tanzania Mozambique Mamonhe 1989IS 4776 India India UP Chari-1 1983 Forage sorghumIS 3923 Zimbabwe Botswana Mahube 1994IS 23496 Ethiopia Tanzania Pato 1995IS 8193 Uganda Rwanda 2001IS 8193 Uganda Kenya Kari Matama 1 2001IS 13444 Zimbabwe Sudan Arous el Rima 2001 Drought tolerantIS 29415 Lesotho Eritrea Shiketi 2000IS 15401 Cameroon Mali Soumalemba 2001IS 21219 Kenya Rwanda 2001IS 25395 Kenya Rwanda 2001IS 33844 India India Parbhani Moti 2002IS 9302 South Africa Ethiopia ESIP 11 1980IS 9323 South Africa Ethiopia ESIP 12 1984IS 30468 Ethiopia India NTJ 2 1980IS 9468 South Africa Mexico Marvilla no SOFO

430201092 2000IS 13809 South Africa Mexico 1990IS 9321 South Africa Mexico 1990IS 9447 South Africa Mexico 1990IS 2391 South Africa Swaziland MRS 13 1989IS 3693 USA Swaziland MRS 94 1989IS 9830 Sudan Sudan Mugawim Buda-2 1991IS 3924 Nigeria India SwarnaIS 3541 Sudan India CS 3541 Converted Zera-zeraIS 3922 × IS 1151 Nigeria, India India 604 Cross progenyIS 3922 × IS 1122 Nigeria, India India 302 Cross progenyIS 2954 × IS 18432 USA, India India 370 Cross progenyIS 2950 × IS 1054 USA, India India R 16 Cross progenyIS 3687 × IS 1151 USA, India India 148/168 Cross progenyIS 6928 Sudan India Moti 1978 Induced mutantIS 18484 × IS 3924 India, USA India SPV 297 1985IS 4283 × IS 18478 India, India India CO-25 1985

Popped sorghum grains are consumed as snack food in parts of India. A screening of 3682selected germplasm accessions resulted in the identification of 36 cultivars with good poppingquality (Prasada Rao et al. 1989).

60

Broomcorn sorghums characterized by long panicle branches are used to make household,warehouse, whisk, toy or hearth brooms (Weibel 1970). These types probably developed in theMediterranean region from material from India or Africa via the Middle East, spread to Italy and thento Spain, France, Austria and southern Germany. The world collection maintained at ICRISAT has 52broomcorn accessions. The spikelets in recently developed cultivars disarticulate before theinflorescences are harvested, eliminating the need for threshing (Prasada Rao et al. 1989).

3.8. Plant QuarantineThe NBPGR, representing the Indian national plant quarantine service, carries out plant quarantinefor all ICRISAT seed and plant material, including sorghum. ICRISAT actively assists NBPGR inprocessing seeds for export, making sure they are free from quarantine objects by using aninspection methodology aimed at ‘exclusion’. Seeds are fumigated and tested by blotter and agarplate as well as by washing.

3.9. ConclusionsThe sovereignty of countries on their own natural resources, affirmed by the CBD and the subsequentGlobal Plan of Action, for conservation and sustainable utilization of plant genetic resources approvedat the International Conference on Plant Genetic Resources in Leipzig, Germany, will be theprinciples guiding ICRISAT’s future genetic resources activities in sorghum. Accordingly, ICRISATwill expend greater research effort in assessing the adequacy of the existing collections, eliminatingredundancy and characterizing the degree of diversity. These assessments will be conducted withspecific countries and will hopefully involve the exchange of unique accessions between ICRISAT andother gene banks. In-country evaluations using descriptors of value to national programs as well asICRISAT, and providing an opportunity to enhance the training of national scientists to assumeresponsibility for their genetic resources will benefit more directly from their use. Future researchwill focus on using in situ conservation of both the crop and its wild relatives in their natural habitats.

Greater utilization of sorghum’s genetic diversity in developing sustainable solutions to basiccrop constraints and in enhancing productivity will be critical in the future. Better access toinformation on collections through SINGER will greatly contribute to this. Efforts will continue tocollect characterization data at ICRISAT or from specific country evaluations, and to consolidateinformation on the collections. The databases will be constantly updated by both ICRISATscientists and other users. The ability to easily and rapidly share information on the potentialbenefit of an accession will lead to greater utilization of sorghum germplasm. These databases willalso be made compatible with other information systems such as GIS and germplasm databases inother regions or plant genome databases with information on sorghum (Eberhart et al. 1997).

Despite the considerable diversity in the germplasm assembled and the identification ofseveral genetic stocks possessing resistance to biotic and abiotic stresses, very little germplasm hasbeen utilized. Probably the Zera-zera landraces from Ethiopia and Sudan were used extensively.The diversity among the five basic races needs to be used to broaden the genetic base in order toproduce improved cultivars for sustainable agriculture. Greater efforts will go into collecting andevaluating wild relatives in view of their potential for insect resistance, particularly shoot fly. In situconservation of both the crop and its wild relatives in their natural habitats will be studied as acomplementary method to ex situ conservation.

61

3.10. ReferencesBandyopadhyay R, Mughogho LK and Mengesha MH. 1988. Sources of resistance to sorghum grain molds.Plant Disease 72:504-508.

Eberhart SA, Bramel-Cox P and Prasada Rao KE. 1997. Preserving genetic resources. Pages 25-41 inProceedings of the International Conference on Genetic Improvement of Sorghum and Pearl Millet, 23-27Sept 1996, Lubbock, Texas, USA.

IBPGR/ICRISAT (International Board for Plant Genetic Resources/International Crops Research Institutefor the Semi-Arid Tropics). 1993. Descriptors for sorghum [Sorghum bicolor (L.) Moench]. Rome, Italy:IBPGR. 38 pp.

ICRISAT (International Crops Research Institute for the Semi-Arid Tropics). 1991. Conserving germplasmfor the semi-arid tropics. ICRISAT Public Awareness Series. Patancheru, AP 502 324, India: ICRISAT.

Mathur PN, Prasada Rao KE, Singh IP, Agrawal RC, Mengesha MH, Sapra RL and Rana RS. 1992.Evaluation of forage sorghum germplasm. Part 2. Pusa Campus, New Delhi, India: National Bureau of PlantGenetic Resources. 296 pp.

Mathur PN, Prasada Rao KE, Thomas TA, Mengesha MH, Sapra RL and Rana RS. 1991. Evaluation of foragesorghum germplasm. Part 1. Pusa Campus, New Delhi, India: National Bureau of Plant Genetic Resources. 269 pp.

Mengesha MH and Prasada Rao KE. 1982. Current situation and future of sorghum germplasm. Pages323-333 in Sorghum in the eighties: proceedings of the International Symposium on Sorghum, 2-7 Nov1981, ICRISAT Center, India. Patancheru, AP 502 324, India: International Crops Research Institute forthe Semi-Arid Tropics.

Murty BR, Arunachalam V and Saxena MBL. 1967. Classification and catalogue of a world collection ofsorghum. Indian Journal of Genetics and Plant Breeding 27:1-194.

Prasada Rao KE and Mengesha MH. 1979. Sorghum and millets germplasm collection in Eastern Sudan.Genetic Resources Progress Report 16. 17 pp.

Prasada Rao KE and Mengesha MH. 1981a. Genetic resources of sorghum from Eastern Sudan. SorghumNewsletter 24:101.

Prasada Rao KE and Mengesha MH. 1981b. A pointed collection of Zera-zera sorghums in the Gambellaarea of Ethiopia. Genetic Resources Progress Report 33. 27 pp.

Prasada Rao KE, Mengesha MH and Gopala Reddy VG. 1989. International use of a sorghum germplasmcollection. Pages 49-67 in The use of plant genetic resources (Brown AHD, Frankel OH, Marshall DR andWilliams JT, eds.). Cambridge, UK: Cambridge University Press.

Prasada Rao KE and Murthy DS. 1979. A Basmati (scented) sorghum from Madhya Pradesh. CurrentScience 48:824-825.

Prasada Rao KE and Ramanatha Rao V. 1995. The use of characterization data in developing a corecollection. Pages 109-115 in Core collection of plant genetic resources (Hodgkin T, Brown AHD, HinthumTJL, and Morales EAV, eds.). London, UK: John Wiley and Sons.

Rockefeller Foundation. 1970. World collection of sorghums: list of pedigrees and origins. New York: IndianAgricultural Program, Rockefeller Foundation. 376 pp.

62

Singh R and Axtel JD. 1973. High lysine mutant gene (hl) that improves protein quality and biological valueof grain sorghum. Crop Science 13:535-539.

Stenhouse JW, Prasada Rao KE, Gopal Reddy V and Appa Rao S. 1997. Sorghum. Pages 292-308 inBiodiversity in trust (Fuccillo D, Sears L and Stapleton P, eds.). Cambridge, UK: Cambridge University Press.

Subramaniam V, Prasada Rao KE, Mengesha MH and Jambunathan R. 1987. Total sugar content insorghum stalks and grains of selected cultivars from the world germplasm collection. Journal of Science, Foodand Agriculture 39: 289-295.

Weibel DE. 1970. Broomcorn. Pages 441-468 in Sorghum production and utilization (Wall JS, and Ross WM,(eds.). Westport, Connecticut, USA: Avi Publishing.