The,lnternatlonal Rice Resel,arc,h Inl,stiitute, Annua,1 Rs'p,o'rl '1968

The International Rice Research Institute Annual Report 1968

LOS BANOS, LAGUNA, PHILIPPINES, 1968 CITY OFFICE: IRRL MANILA HOTEL. MANILA

MAIL ADDRESS: P.O.BOX 583 CABLE ADDRESS: RICEFOUND

The International Rice Research Institute was established jointly by The Rockefeller Foundation and the Ford Foundation in cooperation with the Republic of the Philippines. It was incorporated in 1960 and dedicated in 1962 when the research program began. This annual report, the seventh to be published by the Institute, presents a detailed account of the work done in 1968.

Contents
Board of Trustees. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Personnel. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Director's Introduction " 11 Crop Weather - 1968 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 15 Plant Physiology " 17 Cereal Chemistry " 47 Varietal Improvement " 59 Plant Pathology '.. 77 Soil Chemistry 113 Soil Microbiology 131 Agronomy 147 Entomology 213 Agricultural Engineering 249 Agricultural Economics 307 Statistics 345 Experimental Farm 362 Library and Documentation Center 363 Office of Communication 365 International Activities 375 Training Programs 391 Publications and Seminars 400

Board of Trustees
DR. FORREST F. HILL, Chairman
U.S.A. Consultant, The Ford Foundation.

DR. ROBERT F. CHANDLER,

Director, The International

JR.

Rice Research Institute

DR. RALPH W. CUMMINGS

Associate Director for Agricultural Sciences, The Rockefeller Foundation, U.S.A.

DR. SALA DASANANDA

Director-General, Rice Department, Ministry of Agriculture, Thailand

DR. NOBORU YAMADA

Regional Rice Improvement Officer, FAO, Bangkok

HON. FERNANDO

E. LOPEZ

Vice-President, Republic of the Philippines, and Secretary, Department of Agriculture and Natural Resources

DR. B, P. PAL
Director-General, Indian Council of Agricultural Research, New Delhi

DR.N.

PARTHASARATHY

Formerly Regional Rice Improvement Specialist, FAO, Bangkok

GEN.CARLOSP.ROMULO
President,

University of the Philippines

DR. M. O. GHANI
Vice-Chancellor,

University of Dacca, East Pakistan

DR. JUAN SALCEDO, JR.

Chairman, National Science Development Board, Philippines

DR. GLAUCO P. VIEGAS

Conselho Nacional de Pesquisas, Brazil

Personnel
Admi nistration ROBERT F. CHANDLER, JR., Ph.D., Director A. COLIN McCLUNG, Ph.D., Associate Director DILBAGH S. ATHWAL, Ph. D., Assistant Director JOSE D. DRILON, JR., B.I.M., M.B.A., Executive Officer FAUSTINO M. SALACUP, B.S.B.A., C.P.A., Treasurer IFOR B. SOLIDUM, M.A., Administrative Associate ZOSIMO Q. PIZARRO, LL.B., Administrative Associate PEDRO G. BANZON, LL.B -: Administrative Assistant , Plant Physiology SHOICHI YOSHIDA, D. Agr., Plant Physiologist BENITO S. VERGARA, Ph.D., Associate Plant Physiologist KENJI KURASHIMA, B.S. Age. Chem., Visiting Assistant Scientist" SYLVIA A. NAV ASERO, M.S., Assistant Plant Physiologist FRANCISCO T. PARAO, B.SA, Research Assistant ROBERTO A. LlLlS, B.S.A., Research Assistant EMERITO A. RAMIREZ, B.S.A., Research Assistant EVANGELINE V. MEDINA, B.S. Chem., Research Assistant LORNA G. CASTANEDA, B.S. Chern., Research Assistant EVANGELIT A A. AVELINO, B.S. Chem., Research Assistant Olemistry BIENVENIDO O. JULIANO, Ph.D., Chemist'[] LOURDES J. CRUZ, Ph.D., Assistant Biochemist LYDA C. BAUN, B.S. Chem., Research Assistant GLORIA B. CAGAMPANG, M.S., Research Assistant ARTEMIO V. CARTANO, B.S. Ag. Chern., Research Assistant MIRAFLOR G. CRUZ, B.S. Chern., Research Assistant BERNARDITA V. ESMAMA, B.S. Chem., Research Assistant CYNTHIA C. IGNACIO, B.S. Chern., Research Assistant MYRNA B. NAZARENO, B.S. Chern., Research Assistant EVELYN P. PALMIANO, B.S. Chern., Research Assistant CONSUELO M. PEREZ, M.S., Research Assistant EVEL YN MAE S_TECSON, B.S. Chern., Research Assistant Varietal Improvement HENRY M. BEACHELL, M.S., Plant Breeder TE·TZU CHANG, Ph.D., Geneticist¢ GURDEV S. KHUSH, Ph.D., Plant Breeder CHUKICHI KANDEDA, Visiting Assistant Scientist" RODOLFO C. AQUINO, M.S., Research Assistant LUCILA C. PARIAL, M.S., Research Assistant RIZAL M. HERRERA, B.S.A., Research Assistant JOSE C. DE JESUS, JR., B.S.A., Research Assistant GENOVEVA C. LORESTO, B.S.A., Research Assistant AGAPITO M. GONZALVO, JR., B.S.A., Research Assistant ESPERANZA M. HERNANDEZ, B.S.A., Research Assistant NORMITA M. DE LA CRUZ, B.S.A., Research Aide CARMEN M. PAULE, B.s.A., Research Aide REYNALDO L. VILLAREAL, B.S.A., Research Aide ESPIRIDION T. TORRES, B.S.A., Research Aide ALICIA O. ALCALA, B.S.A., Research Aide

Plant Pathology SHU·HUANG OU, Ph.D., Plant Pathologist KEH CHI LING, Ph.D., Associate Plant Pathologist CELESTINO T. RIVERA, M.S., Assistant Virologist FAUSTO L. NUQUE, M.S., Assistant Plant Pathologist PRISCILLA C. SANCHEZ, M.S., Research Assistant JOSE M. BANDONG, M.S., Research Assistant TORIBIO T. EBRON, JR., B.S·.A., Research Assistant MANUEL K. PALOMAR, B.S.A., Research Assistant JOSELlTO P. SILVA, B.S.A., Research Assistant SILVINO O. MERCA, B.S.A., Research Assistant VLADIMARTE M. AGUIERO, B.S.A., Research Aide SONIA P. EBRON, B.S.M. Tech., Laboratory Aide Soil Chemistry FELIX N. PONNAMPERUMA, Ph.D., Soil ChemisttP RUBY U. CASTRO, M.S., Assistant Chemist TERESITA A. LOY, B.S. Chem., Research Assistant ELSA R. GUEV ARRA, B.S. Chem., Research Assistant ZENAIDA M. CANLAS, B.S. Chern., Research Assistant CARMEN M. VALENCIA, M.S., Research Assistant MYRNA R. REALUYO, B.S.A., Research Assistant DOMINGO C. RIEGO, B.S.A., Research Assistant SYLVIA G. SAMANIEGO, B.S. Biochem., Research Assistant Soil Microbiology TOMIO YOSHIDA, Ph.D., Soil Microbiologist ELLEN M. BAUTISTA, M.S., Assistant Soil Microbiologist ROSABEL R. ANCAJAS, B.S.A., Research Assistant TERESITA F. CASTRO, B.S. Chern., Research Assistant BENJAMIN C. PADRE, JR., B.S.A., Research Aide Agronomy SURAJIT K. DE DATTA, Ph.D., Associate Agronomist DONALD E. SEAMAN, Ph.D., Visiting Weed Scientist'" DUANE S. MIKKELSEN, Ph.D., Visiting Scientist** HELLENIUS ten HAVE, Ph.D., Visiting Scientist] DONALD L. PLUCKNETT, Ph.D., Visiting Scientist'[ EMMANUEL T. FLORESCA, M.S., Assistant Agronomist ROBERTO T. BANTILAN, B.S.A., Research Assistant PABLO M. ZARATE, B.S.A., Research Assistant CARMELITA P. MAGNAYE, B.S. Chern., Research Assistant JESUS T. MAGBANUA, B.S.A., Research Assistant SERVILLANO N. BALAOING, B.S.A., Research Assistant WILMA N. OBCEMEA, B.S. Chern., Research Aide PAUL C. BERNASOR, B.S.A., Research Aide VICENTE C. FALCULAN, B.S.A., Research Aide RENE Q. LACSINA, B.S.A., Research Aide Multiple Cropping RICHARD B. BRADFIELD, Ph.D., Agronomist FRANCISCO L. VALBUENA, B.S.A., Assistant Agronomist WILHELMINO T. HERRERA, B.S.A., Research Assistant Entomology MANO D. PATHAK, Ph.D., Entomologist FAUSTO L. ANDRES, B.S.A., Research Assistant GERARDO B. AQUINO, B.S.A., Research Assistant JOSE I. CALDERON, B.S.A., Research Assistant DENIS T. ENCARNACION, B.S.A., Research Assistant SUSAN R. HIZON, B.S. Chern., Research Assistant

EDWIN S. RAROS, B.S.A., Research Assistant NAPOLEON C. RESPICIO, M.S., Research Assistant .CARLOS R. VEGA, B.S.A., Research Assistant

Agricultural Engineering
AMIR U. KHAN, Ph.D., Agricultural Engineer LOYD H. JOHNSON, M.S., Agricultural Engineer=" STANLEY S. JOHNSON, Ph.D., Agricultural Economist*'" DW A YN E A. SUTER, M. S., Consulting Agricultural Engineer'[ EMILIO O. CASEM, M.S., Assistant Agricultural Engineer MAKOTO HOKI, M.S., Visiting Assistant Design Engineer ANTERO S. MANALO, B.S.A., Research Assistant NORBERTO L ORCINO, B.S., Research Assistant BEN HUR AGUILA, B.S.A., Research Assistant ELiSEO L. RUIZ, B.S.A., Research Aide ABDUL I. AMILHUSSIN, B.S.A.E., Research Aide BENIGNO T. SAMSON, B.S.A., Research Aide NESTOR C. NA VASERO, B.S. A., Research Aide FELICI ANO C. J ALOTJOT, Draftsman FERNANDO S. CABRALES, B.S.M.E., Draftsman

FLORENCIO F. MACAPUGA Y, B,S. Ag. Adm., Rice Production Assistant ANDRES V. FORONDA, B.S.A., Rice Production Assistant SATURNINO E. RONQUIllO, B.S.A., Rice Information Assistant LEONARDO T. ALMAZAN, Research Aide ARNULFO C. DEL ROSARIO, B.SA, Artist-Illustrator FEDERICO M. GATMAITAN, JR., Artist-Illustrator EDITO S. RUFON, MultiJith Operator CECILIO L. PALACPAC, JR., Assistant Multilith Operator URBITO T. ONGLEO, B.S.A., Photographer FELICIANO J. TOYHACAO, Assistant Photographer ANTONIO O. VALENCIA, Assistant Photographer

Library and Documentation

Center

Agricultural Economics
RANDOLPH BARKER, Ph.D., VIOLET A G. CORDOVA, B.S. HILDA G. CURA, A.B. Econ., CRISTINA M. CRISOSTOMO, Agricultural Economist A., Research Assistant Research Assistant A.B. Econ., Research

LINA MANALO-VERGARA, M.S., Chief Librarian GLORIA S. QUIROS, A.B., B.S.L.S., Assistant Librarian MILAGROS C. ZAMORA, M.S., Bibliographer ADEL1NA D. FERRER, B.S.E., Indexer CARMELITA S. AUSTRIA, B.LS., Circulation Librarian MILA C. MEDINA, B.L.S., Order Librarian GREGORIO A. ARDALES, Binder JUKYU CHO, D. Agr., Translator (in Japan) ETSUKO TAKEYOSHI, B.A., Indexer (in Japan) TAEKO ISODA, B.A., Indexer (in Japan)

Assistnat
ERIBERTO ABRAHAM

R. ACUNA, B.SA, Research Aide M. MANDAC, Research Aide

Food and Dormitory

Services

Statistics
KW ANCH AI A. GOMEZ. Ph.D., Associate Statistician ROSALINDA C. ALiCBUSAN, B.S.A., Research Assistant FLORENCIO A. ARICA, JR., B.S. Econ., Research Assistant WILFREDO C. GOMEZ, B.S. Chern. Engr., Statistical Aide EMERITO V. TIPA, E.S.A., Statistical Aide

Experimental Farm
FEDERICO V. RAMOS, M.S., Associate Agronomist and Farm Superintendent+ ORLANDO G. SANTOS, B.SA, Associate Farm Superintendent JUAN M. LAPIS, B.S.A., Research Assistant ELADIO M. BARADAS, B.S.A., Research Aide FILOMENO 0. LANTING, B.S.A., Research Aide

REBECCA C. PASCUAL, M.S.H.E., M.S., Manager NENITA C. ESGUERRA, E.S.H.E., Assistant Manager ESTER P. NOVERO, B.S. Home Tech., Head Food Supervisor ESPERANZA A. ENRIQUEZ, Matron AURORA T. VERGARA, B.S. Home Tech., Assistant Matron PATRIA E. MENDOZA, B.S. Home Tech., Food Supervisor ESTELA G. D1VINAGRACIA, B.S. Nutrition, Food Supervisor ELISA C. ESTRADA, B.S.F.N., Food Supervisor VIOLETA C. MERCADO, B.S.F.N., Food Supervisor NENTA E. QUIJANO, B.S.F.N., Food Supervisor

Buildings and Grounds

HERMENEGILDO G. NAVARRO, B.S.M.E., Property Superintendent VICTOR T. ARANEZ, Assistant to the Property Superintendent RIZALINO T. DILAG, JR., B.S.A., Grounds Supervisor

Office of Communication
WILLIAM G. GOLDEN, JR., M.S., Rice Production Specialist MERRY LEE C. SAN LUIS, A.B., Editor" ROGELIO D. FELICIANO, M.A., Associate Editor FRANCISCO D. GORREZ, JR., B.S.A., Rice Production Technician INOCENCIO C. BOLO, B.S.A., Rice Production Technician RAMIR9 C. CABRERA, B.F.A., Graphic Designer CORAZON V. MENDOZA, Litt. B., Editorial Assistant EUSTACIO U. RAMIREZ. B.S.A., Rice Production Assistant ROMARICO S. NECESARIO, B.S.A., Rice Production Assistant

• Arrived during year. •• Left during year. t Arrived and [eft during year. tt On stu dy leave March to Dec., 1'168. IjJ On study leave since Aug., 1'168. + On study leave since July, 19.68.

Director's Introduction
During 1968, the Institute made a strong effort to identify new genetic lines of rice from its breeding program that would have all the advantages of IR8 and IRS and none of their disadvantages. Although none of the selections were pure enough or had been sufficiently tested to allow them to be named in 1968, enough promising materials were identified to give us confidence that one or two new varieties will be named in .1 969. Each year the Institute staff tests more than 40,000 genetic lines from many different crosses. In fact, since the Institu te started its research program more than 1,500 crosses have been made. It is only a matter of time before outstanding varieties will be developed and identified which have the high yield potential and fertilizer responsiveness of IR8, but with superior grain quality and increased resistance to the major diseases and insect pests. Among other things being developed by the Varietal Improvement Department are varieties adaptable to deep water, high-yielding Basrnati types, and earlymaturing varieties (100 days from seed to seed). The outstanding contribution of the Agronomy Department during 1968 was to identify and test several promising herbicides, or combinations of herbicides, that would effectively control weeds in transplanted rice. Trifluralin and MCPA; and o:-2,2,2-trichloroethyl styrene and the isopropyl ester of 2,4-0; applied once every 4 days after transplanting gave highly satisfactory control of weeds, grasses, sedges and various broad-leaved species. All three of these chemicals, including the MCPA or 2,4-0, can be applied in a granular form. This eliminates the problems connected with purchasing and maintaining spraying equipment. Also, these chemicals are less costly than handweeding, and two of the three chemicals mentioned are already on the market in the Philippines and hence available to farmers. The Agronomy Department continued to work with upland rice. In 1967 excellent yields were obtained, but in 1968 the rainfall was quite uneven and lower in total amount. As a result yields were extremely low. This simply shows that rice has low tolerance to drough t and in regions where rainfall is highly unpredictable, it would be preferable under upland conditions to plant crops other than rice, such as sorghum, millet, soybeans and maize. The Agronomy Department has been attempting to increase total annual rice production by growing four crops a year. During 1968, four crops of the variety Taichung (Native) I yielded a total of 22,400 kg/ha. This is the highest total annual production of rice yet obtained at the Institute. An early-maturing selection from the IR8 line produced 22,000 kg/ha and another variety which we call "dwarf Peta" yielded 21,800 kilograms. All of these yields were obtained in 365 days or less. The Agronomy Department continues to conduct a few experiments on farmers' fields and in one in Calamba, IR8 yielded 10,474 kg/ha. This is the highest yield yet obtained on replicated plots in the Philippines. Every effort was made to keep the cash inputs as low as possible, and this high yield was obtained by spending a total of 1"zI.s6 for insect and weed control, and for fertilizers. This is of course the cash outlay and does not involve the cost of labor, other than that of weeding. Ten tons of rice should sell for approximately

n,300. It would indeed give a farmer a good

11

labor income from only one crop, and of course, if he has irrigation water, he can grow one or two additional crops during the year. The most striking contribution of the Entomology Departmen t during 1968 was the fu rther proof that a high degree of immunity to plan thopper and leafhopper attack exists in several varieties from India and that this quality is rather simply inherited and can be readily transmi tted to improved bu I susceptible varieties. For example, the variety Mudgo from India is resistant to the brown planthopper, which causes extensive direct damage and is also the vector of the grassy stunt virus disease of rice. Planthoppers caged wi th this variety live no more than 4 days and do no t mu It ipl Y su ffic ie n t Iy to rna in ta in the ir popu Ia ti on. When Mudgo is crossed with IRS, which is a highly susceptible variety, the progeny in the F2 generation show a 3: 1 segregation, that is, there are three resistant plants to one susceptible. Although the genetics of this have not yet been carefully studied, it appears to be a straigh I Men de lian ra tio indica ling that a single domi n an t ge ne is responsible for the resistance. If there are modifying genes, their influence must be sligh t. A similar study is being carried OUI with the green planthopper, also a vector of virus diseases. A variety called Pankhari 203 from India has strong resistance to the green leafhopper. The resistance can be incorporated into a susceptible variety in a sim il a r way as is the resistance to the brown pi an t hoppe r. Prelim ina ry ev idence indica tes that soon we shall to able to develop improved varieties that have strong resistance to both the brown plan thopper and the green leafhopper. This will indeed be a great boon to farmers who now must either spend large amounts of money for insecticides to can trol these pests or must suffer the yield losses that result from their attacks. There was widespread evidence obtained this year to indicate that the brown plan th oppe rs on the Insti tu te 's fiel d we re begin n ing to develop re sistance to diazinon, the chemical which originally gave good control. The Institute has been using this chemical for about 3 years and with a new generation of brown planthoppers every IS to 20 days, obviously many generations have occurred. We are now testing several new chemicals for insect control on rice and some of them look highly promising. The Chemistry Departmen t con tinued to study the protein content of rice grain and the distribution of the essential amino acids. It was mentioned in the 1967 report that quite a few varieties averaged more than 13.5 percent protein in the brown rice (the average protein content of brown rice is about 8 percent). Several of these selections have been crossed with varieties with a good plant type, such as IR8. Very preliminary information indicates that there is some possibility that high-yielding, nitrogen-responsive varieties of rice can be developed that have protein contents averaging \0 percent rather than less than 8 percent The protein conten t of rice is strongly influenced by environment and, generally speaking, as yield goes up the protein content goes down. It may take another 2 years to determine for certain whether a significant and consistent high protein level can be in trod uce d in to rice var ieties through pi an t b ree ding. As mentioned in the 1967 report, the Agricultural Engineering Department has developed a new type of mechanical thresher. During 1968 this piece of equipment was further improved and widely tested. It has proved to be quite successful from the standpoint of acceptance by selected farmers, principally because it is simple to operate, has few mechanical troubles, and can thresh wet rice. Three commercial companies in the Philippines have constructed models after the Institute design, and

12

some of these manufactured models are now being shipped to other cooperating cou n tries fo r trial. The Engineering Department continues to work on equipment for wet land preparation, but it may be several years before something significant will come from the program. Work was started in 1968 on a mechanical stripper for harvesting rice. The Multiple Cropping program of the Institute was expanded during 1968 because of the increasing awareness of the need and opportunity for growing food crops the year round in tropical Asia. Essentially all the land in the Multiple Cropping experimental area was plan ted to rice during the monsoon season and to several other crops during the remainder of the year. Among the more successful crops in rotation with rice are soybeans, sweet potatoes, sweet corn, cow peas and sorghum. The program received a grant from The Rockefeller Foundation enabling the construction of a small building for multiple cropping which will have a classroom for training, offices, storerooms, and workrooms for the staff and trainees. It is expected that a mu Itiple cropping training program for the Philippines and Southeast Asia will be underway by June, 1969. The Rice Production Training Program continued unabated and at an international level. A group of 35 scholars from nine countries were in residence at the Institute from early June until early December, 1968. The largest group was from India; other countries represented included Pakistan, Ceylon, South Vietnam, Indonesia, the United States, Malaysia, Laos, and the Philippines. The spread of Institute varieties continued throughout Asia and in certain other parts of the world. For example. Pakistan is now growing more than one million acres of I R8 and India may have a total of five million acres. Burma has an estimated 400,000 acres and the Philippines has more than one million acres. Undoubtedly, in the world there were at least 10 million acres of IRS being grown in 1968, and if you estimate conservatively an additional profit of U.S. $60 per acre, the calculated increased income for rice farmers in 1968 is $600,000,000. During 1968 the expenditures of the Institute was somewhat over I \tl million dollars which shows that there is a rather high pay-off on the investment made by the foundations and other organizations supporting the research program.

Staff Changes
Mrs. Merry Lee San Luis was appointed as Editor of the Institute replacing Mr. Edward A. Jackson who resigned in 1967. on July 1,1968,

Trustees
Dr. Noboru Yamada, the regional rice information specialist of the Food and Agricultu re Organization in Bangkok, was elected a trustee of the Institute for the 4-year period beginning January 1,1968. Dr. Yoshiaki Ishizuka's 4'year term expired on December 31, 1967.

Finances
As has been true previously. the major portion of the operating costs of the Institute was shared equally by the Ford and Rockefeller foundations. The sum rece ived during 1968 from the two foundations for the lnstitu te 's cen tral program amounted to $1,544,550, which included 5300,000 provided by The Rockefeller Foundation in terms of manpower services and their travelling expenses.

13

Other funds received by the Institute in 1968 are listed below: 1. From the Ford Foundation for support of rice research and for the training of scientists and extension West Pakistan Ceylon Philippines workers. as follows:
$ 80.000

147,660 40,890

The Foundation also provided $80,000 to support the training and cooperative research program of the lnstitu teo 2. From The Rockefeller Foundation, to support the training and cooperative program of the Institute, $93,000, and to support the accelerated research and training program on cropping systems for 'tropical areas, $76,800. 3. From the United States Agency for International Development, a total of $104,216.99. Of this amount $54,677.34 was for support of a research project entitled "Research on Farm and Equipment Power Requirements for Production of Rice and Associated Food Crops in the Far East and South Asia"; $32,494.65 was for the accelera lion of the rice research program in India; and $17,045 was a contribution toward the training program of the Institute. Furthermore, the Agency for International Development has given the Institute a $400,000 grant toward the cost of expanding the Institute's training and consultation services; the grant was made available for the period july I, 1968 to June 30, 1969. 4. From the National Institutes of Health, $65,127.25 for the screening of rice varieties with respect to their amino acid content, particularly as regards lysine and threonine, and to study the genetic characteristics of the lysine-rich varieties. 5. From the National Science Foundation, $5,750 to support a research project entitled "The Description and Preservation of the World's Rice Gerrnplasrn, " 6. From the National Science Development Board, :P44,493 ($11,408.46), to support the studies on the "Insecticide Residue Problems of Rice Production with Emphasis on Gamma-BHC." 7. From the Imperial Chemical Industries, Ltd., $10,000 to support research on minimal tillage techniques, and on the use of non-selective herbicides for tropical rice production. 8. From the Stauffer Chemical Company, $5,000 as its contribution toward the general research program of the Institute. 9. From Union Carbide Asia, Ltd., $5,000 to assist the rapid dissemination of the Institute's research findings, particularly in the field of entomology. 10. From the J. R. Geigy S. A., $5,000 to partially support the insecticide program of the Institu teo 11. From the Esso Research and Engineering Co., $4,305.92, a contribution toward the fertilizer research work of the Institute. 12. From the International Potash Institute, $2,400 as its contribution toward the soil fertility studies of the Institu teo 13. From the Eli Lilly and Co. (ELANCO), $2,000 to partially support the Institu te's weed control research program. 14. The Bio-Products Dow Chemical International, $1,537.30 to help support the weed and grass control research of the Institute.

14

Crop Weather -1968

good grain yields. The amount of solar energy received during the critical period of the 1968 wet1959 - 1967 A.e(QQe season crop (September to November) - when most of the varieties were in the grain development ~ W~------------..-~~~~-~--4~-_,,~.--_, stage - was higher than that received during the same months in 1967. The other significant aspect of the 1968 crop weather was the low frequency of typhoons during the wet season. The first typhoon, "Didang", which occurred on July 23-25 with a maximum wind velocity of 61 km/hr did not cause damage to the rice because most of o a '2 16 <0 Z4 ae 'Z '640 44 48 52 FMAMJ J A SOND the crop was newly transplanted. Two other minor Months typhoons, "Huaning" and "Nitang", occurred on Fig. I. Weekly rainfall curve (3-point moving average) for August 18-20 and September 27-29, respectively, College, Los Banos, Laguna. both with a maximum velocity of only 40 km/hr. The low frequency of typhoons and slightly higher solar energy received during the ripening period of the 1968 wet-season crop resulted in high grain yields varying from 6,000 to 7,000 kg/ha in many The weather during the 1968 dry season (January through May) was normal compared to recent of the experiments.
100,----------------------------------,

averages. During the wet season (June through November) there were a few deviations from the normal. Figures 1 and 2 show the curves for rainfall and solar radiation recorded by the Philippine Weather Bureau station at College, Laguna, located about 700 meters from the Institute farm. From January to November 30, 1968, the rainfall amounted to 1,263 mm as compared to 1,769 mm for the 19-year mean (1959-1967). At the beginning of the wet season, at which time the land is usually being prepared, the rainfall was normal compared to the long-term average. The abnormally low rainfall during the monsoon season greatly reduced the yields of rainfed, upland rice (nonflooded). In fact, the yield data were so variable that they are not recorded in this report. The total solar energy received during the ripening period (April and May) of the 1968 dry season crop was 1,411 g-cal cm-2 lower than that of 1967 and 2,973 g-cal cm-210wer than the longterm average. This had no appreciable significance, however, and the high-yielding varieties produced

3500,-----1959-1967 A.erage

e
c
<>

s
'i:i

'"

2500

a:

l;i
0 (fl

2000

4 6 12 15 JFMAMJJASO

16 24

25 32

36 40

44 48 ND

52

Months

Pig. 2. Solar radiation curve (3-point College, Los Banos, Laguna.

moving average)

for

'The general climatic and soil environment at the III·· stitu te was described in the 1965 Annual Report and the crop weather for 1966 and 1967 were described in the annual reports for those years.

Crop Weather-1968

15

Plant Physiology
Studies on the varieties of improved plant type showed that their physiology differs from that of the traditional tropical indica varieties. The improved variety fR8 has a steady high growth rate from the seedling stage to harvest whereas the traditional variety Peta has a high growth rate at the early stages followed by a low growth rate after flowering, resulting in low yields. IR8 can produce more dry matter than Peta with the same leaf area index, and has a larger optimum leaf area index and a higher maximum crop growth rate. It seldom exceeds its optimum leaf area index: consequently, it can respond to high levels of nitrogen without showing a noticeable decline in grain yield. These results explain some of the differences in the response to nitrogen of IR8 and Peta. Detailed analysis of the yield components of improved varieties indicated that the number of grains per unit area is an adequate measure for evaluating growth performance. The model yield components of transplanted IR8 are given in this report. Detailed studies were also made on the tillering performance of rice varieties as affected by nitrogen nutrition. Studies on photosynthesis showed a higher photosynthetic rate for the varieties tested than those reported so far in the literature. The experiments on photoperiod showed that long photoperiods after panicle initiation can delay panicle exsertion, a delay mainly in development and elongation. Temperatures between 21 and 32 C during photoinductive cycles did not affect panicle initiation. Salinity studies dealt with the different kinds of the salt and their effect on potassium supply. It appears that the antagonism between sodium and potassium in the rice plant occurs only when the potassium supply is relatively high . Work on zinc deficiency centered on factors affecting zinc uptake and on correcting zinc deficiency. Dipping of rice seedlings in zinc oxide suspension was found to provide an effective and economical way of correcting zinc deficiency.

., '"
'0.
<:

.9-

..c: f-

.,

...:
a:
D..

o
w a:
_l

=> z z

«
QO

CD In

17

Growth Rate and Plant Characters of Rice Varieties in Relation to their Response to Nitrogen
Previous studies of the growth rate of both the japonica and the leafy, tropical indica varieties of rice have shown that a high growth rate at the early stages is followed by a low growth rate at the later stages, and vice-versa. Since most of the grain carbohydrates are actually photosynthesized after flowering, varieties with a high dry matter production during the later stages generally give high yields. It has been established that the high growth rate of the tall, leafy, tropical indica varieties in the early stages tends to result in excessive vegetative growth, which in tum causes mutual shading and subsequently a lower growth rate. On the other hand, rapid seedling growth is considered a desirable trait of tropical rice varieties, particularly when combined with early maturity. It enables the young plants to become fully established before weeds become a problem. Obviously, too, it is associated with heavy tillering capacity. The new dwarfvariety, IRS, with its stiff straw and short, erect leaves, offers a good opportunity for studying the relationship between the growth rate at the different physiological stages and the yield of the new tropical varieties now being developed. As the studies reported in this section will show, the modern plant types appear to behave quite differently from the traditional tropical indica varieties.
Seedling growth and grain weight

10 Sand:

10-Day Experiment
00<1<

• •• ••

• ,•

••
Soil: 00rI<

,
li9h1

:•• • ••
•

•• •

So ... :

Soil: UQI>!

10

• •••
17 19 21

•I ••• • ••
23 25

•• ••
17

•• •• •• • • •
21 23 25

0'5

'Z7 Z9 IOOO-Grain

15

19

2.1 29

Weight (9)

Fig. I. Relationship between shoot weight and grain weight under various conditions.

Pre-germinated seeds of 14 varieties with different grain weights were grown in the greenhouse and in a darkroom for 10 and 20 days. As shown in Fig. I, the dry weight of shoots in the IO-day experiment seems to be well correlated with grain weight: the heavier the grain weight, the larger the shoot weight. This suggests that early growth rate in rice Varieties depends largely on the reserve nutrients in the seeds. A comparison of seedling growth in the light vs. dark and clay vs. sand treatments indicated that (a) the seedlings grown in the light had a faster growth rate than those grown in the dark and (b) the clay-grown seedlings had a faster growth rate than the quartz-grown seedlings only in the light. This

indicates that nutrient supply may have an effect on growth only in conjunction with photosynthesis. In the 20-day experiment, the relationship between shoot weight and grain weight was less exact, but there was a close relationship between shoot weight and leaf blade weight (Fig. 2). This suggests that leafiness becomes the dominant factor at the later stages of seedling growth. Plant height was well correlated with shoot weight (Fig. 3). However, seedling height was not necessarily correlated with plant height at harvest, there being a large variation in plant height at harvest among seedlings of similar sizes (Fig. 4). This indicates that rapid seedling growth is not necessarily associated with the tall, leafy varieties at later stages. Early vegetative growth and LAI IR8 and Peta were grown on the Institute farm at 30 x 30 ern and 100 x 100 em spacing and in combination with three levels of added nitrogen. The plants of the Peta population were supported with bamboo stakes whenever there was danger of lodging. As shown in Fig. 5, Peta is leafier than IRS. At the early growth stages when the leaf area index (LAI) is small, the growth rate of the crop was almost a linear function of the LAI, and was

IS

Plant Physiology

400

...
l-

20- Day Experiment

•
I I ..

.s
0

~320

l=

240

l-

• •
•
....

• • •• • •
I

~ tJ)

~ 160 120 15

••
60

••
I

•••

•
I

••

•• ••

19 23 27 l()()()-Groin Weight (0)

140 180 100 Leaf blade Weight (rng)

220

Fig. 2. Relationship of shoot weight to grain weight and to leaf blade weight.

70

200

•
60 r180

••
-

E :=.

50

I-

...E

• •• • •

.3
0

01

! 40 r-

•• ••• • •• ••
I

"1n 160 ~

15
L;

'i» 140 ~

01

30

120

l-

20 100

200

_L 300

400

100 30

•• ,
40

..

•
I

50

60

Shoot Weight (mg)

Fig. 3. Relationship between shoot weight and height of 2O-day-old seedlings.

Height of 20 day-old Seedlings (em)

Fig. 4. Relationship between plant height of 2~ay-old seedlings and at harvest.

Plant Physiology

19

Table I. Photosynthetic rate of isolated leaves of IRS and Peta at low and high light intensities.

140.

120

VarietY

Nitrogen level (kg/ha)

mgC0211

Photosynthetic rate OOcm2/hr

10 KLux 60 KLux

N content of leaf blade

(%1

IRe

0. PeT.

IR8

0 100 (mean)

18.2 21.1 19.7 23.1 23.4 23.3

44.1 49.8 47.0 4S.5 43.8 46.7

4.16 6.24 5.20 4.63 5.28 4.96

40

Peta

0
100 (mean)

20

v1V °
0.2

/•

/.

/ /.

V
0

4 -6 _

aile< Iransplanli"IJ

0.6

1.0

1.4

1.9

22

2.6

Mean LAI

independent of variety (Fig. 6). At the 100 x 100 em spacing, a close relationship between LAI and growth rate existed at all growth stages, largely because of the reduced mutual shading at the wide spacing. Photosynthetic rate of isolated leaves The photosynthetic activity of a population (P I) per unit land area is a product of the leaf area index (A), the light-receiving coefficient (f), and the photosynthetic rate per unit leaf area under
1.6 _ 1.4 1.2 1.0. 0..8 0..7 0..6 0..5 0..4

Fig. 6. Leaf area index and crop growth rate.

E :r

Ii"

~o..8 0..6

&0.4 0..2 0. 8 7 6

Q2

0..1 0. 4.0. 3.S 3.0.

~.~

001, 00

an

<i
..J

5 4

ZD
1.5 1.0.

3
I 0. 0.246810121416

ligh t -saturated condition (Po). Thus, PI =Ax f x P, The leaf area index (A) can be changed by such cultural practices as increasing the plant density and fertilization; the light-receiving coefficient (f) is affected by LAI and such varietal characters as leaf erectness; and the photosynthetic rate per unit leaf area is controlled by varietal characters and nutrition at a given stage. To find out what factor (s) is dominant in determining PI, it is necessary to separate Po from A and f. The photosynthetic rate of IR8 and Peta per unit leaf area are about the same (Table 1). This implies that when the PI'S of the two varieties are to be compared, Po is no longer a determining factor, and thus PI is determined by A and by morphological characters. This conclusion offers a basis for evaluating plant type in the determination of'P, or dry matter production. The photosynthetic rate of leaves is affected by nitrogen nutrition. The higher the nitrogen content, the higher the photosynthetic rate (see Fig. 14). Plant type, optimum LA', and maximum eGR Up to 8 weeks after transplanting, the dry weight of Peta was heavier than that of IR8 at the 30 x 30 cm spacing; at later growth stages, this situation

0.

t-_"'---'--'-.L...J__.l___j
246810121416

Weeks 011 .. TltlospIan~1I9

Fig. 5. Dry weight and LAI of IRS and Peta at successive stages of growth (100 kg/ha N).

20

Plant Physiology

Table 2. Crop growth rate (CGR), mean leaf area index (LAI), and nitrogen content of the leaf blade of IR8 and Peta at different nitrogen levels and spacings, IRRI, 1966 wet season. 4th·6th week" Variety Nitrogen Spacing applied (em) (kg/ha) CGR** Meant LAI NARtt N%+ CGR 8th· 10th week * Mean LAI NAR N%

IR8

30 x 30

0 50 100 0 50 100 0 50 100 0 50 100

49 71 92 74 104 127 13 14 15 14 20 16

1.15 1.60 1.91 1.37 2.26 2.56 0.18 0.20 0.23 0.20 0.26 0.35

43 44 48 54 46 50 72 70 65 70 77 46

3.70 3.85 4.40 3.34 3.60 3.87 4.00 4.29 4_66 3.64 3.68 4.00

118 136 191 105 127 137 36 43 47 50 47 60

2.93 3.14 5.76 3.38 4.72 5.76 0_69 0.79 1.00 0.98 1.25 1.41

40 43 33 31 27 24 52 54 47 51 38 43

2.39 2.50 2.68 2.22 2.28 2.36 3.38 3.56 3,78 2.96 2.78 3.14

Peta

30 x 3G

IR8

100 x 100

Peta

100 x 100

• After transplanting. * * g/sq m/week. t Mean of LAI 's at the start and end of the 2-week period. tt Net assim ilation rate. + Nitrogen content of leaf blades.

was reversed (Fig. 5). However, the LAI of Peta was always larger than that of IR8 and reached about 8 at flowering. This indicates that mutual shading in the Peta population became serious and this reduced the efficiency of dry matter production per unit leaf area. To study the quantitative relationship between plant type and optimum LAI, data were collected from the population 8 and 10 weeks after transplanting. Crop growth rate (CGR) is defined as the increase in dry matter in grams per square meter per week, and net assimilation rate (NAR) as:

NAR =

CGR~(~Y_:_) ~
Mean LA! (F)

Y F

(1)

Therefore, the NAR is considered as a measure of leaf efficiency for dry matter production. From Table 2 and Fig. 7, it is obvious that the NAR decreased with the increase in mean LAI. In other words, the NAR is dependent on the mean LA! which determines the average light intensity in the leaf layers of a population. It is also logical to assume that the higher NAR values for the 100 x 100 em spacing than for the 30 x 30 cm spacing are partly due to higher nitrogen content because the photosynthetic rate of a leaf is a function of

Plant Physioiogy

21

Table 3. Some plant characters of IRS and Peta at flowering (100 kg/ha N), 1966 wet season.

60

Variety IRB Peta

Height (em)
121

Mean leaf openness

22.1 34.2

Leaf blade ratio * 0.56 0.46 ratio

50

K""
40

<,
~
0

...............
......

0.40 0.51 (in

~ 30

~ <, <, ~

196

~ ...... ~

Leaf blade to leaf-sheath-plus-culm weight). ** Ught extinction coefficient weeks after transplanting.

..

r-.......... ......

.IR8

20
8-10 10

o Peto
_ks ofter tron,planting

i'z

--,

measured at 10

3 Mean LAI

its nitrogen content, other factors not being limiting. Thus, the NAR measured in this experiment is considered a function of the mean LAl and the nitrogen conten t of the leaves. Nevertheless, as shown in Fig. 7, there is linear relationship between NAR and mean LAI (F):
NAR ==a-b F

Pig. 7. Relationship between net assimilation and mean leaf area index (LAIl.

rate (NAR)

(2)
(3)

..
.,.
E

180

_IRS

o Pete

140

By rewriting equation (1):

Y ==NAR

2100

....

xF

From equations (2) and (3), the following parabolic equation between CGR and mean LA! was obtained:
Y ==a F - b F2 (4)

60

20

• Meon LAI

'"

By this mathematical manipulation, it seems justified to fit data on CGR and LAl into a parabolic equation with the help of the least square method. As shown in Fig. 8, the relationship between mean LAIand CGR can be expressed by a parabolic equation.
IR8 Peta : Y==52.1 F- 3.29 F2
(5)

Fig. 8. Op tirnurn leaf area indices of IR8 and Peta,

Opt. LAI IR8 Peta

Max. CGR

7.91
5.94

206 g/sq.lm/week 135

: Y ==45.3 F - 3.81 F2

(6)

The optimum tion:

~==O dF

LAI was obtained by the equa-

Substituting the optimum LAI into equation (5) and (6) gave the maximum CGR for each variety. The following data summarize the above calculations.

These results show that IR8 can produce more dry matter with the same LA!, and has a larger optimum LAI and a larger maximum CGR. These results explain also why IR8 and Peta differ in their response to nitrogen. With increasing levels of nitrogen, Peta tends to exceed its optimum LAl, resulting in a decrease in the amount of dry matter it produces. On the other hand, IR8 seldom exceeds or, when it does, only to a small extent, its optimum LAl. Con-

22

Plant Physiology

iequenUy ,it C<:!1'l [es,poorldto 11,it?JluH ,Icve~s of niUQs~nw,i[hcut ShQwi.!il.g~,~QUi;c:!!.b]e decii:Jl1cm g~ah~ y[~I,d.. Thi~: a~'~a!'SlQ be the ph;y~iQI gitil o 'iJas,jsof Ole ,mtmg,en rospo!l1.siv(!m:ss of :~ R8. S][loe theJi'l ~IDO ,~igtlmc:!!_!Jlt diffe~re!1ce Ln the pho~(!sYl1Jlhiilti,crn~e of isolated .~,~'av(!'s !llW"Ge~ th~ b two \'!ll!'i~,t~e~, C;;I1'l be condudc,d UI~! HI,C' i:n~ H c.re:l~i'ld opitw!l:J!IIUn tAl of lR8; ii~ the ovendJ [e~111 t of ~h:eiml,)mVeln~n~; :nn plan ~ type. T,i~ewfOlrc, oomne p]@!1[ chru;"'ctenof the two varie:tics that .-we Jcl,!I:~'c!~O d:ry ImUe~ p.ro(]lIct.ioJil. W~roiIlVC-Sli·
gated.

('1.orn.tl1l.1siron

nti~iro:.pllrimellt.

IIppe~tQit!dicat~

that t&'tc

do:milllilant (a.clef wtLich deter.mill[lesthe~ow& r.OI.w of wiice varn:eMes vatriiesMth lilie, .gfQwth stage. At

lhe ~dl:mt na:~,e:. the ;~weight ]aQe~Y mfJIUIef.!~~~lhepowtilit rate;. :m;I.b.!i'C!qjlJtef!~ly, ru~aJfm~SIl 'Of h:ilif i.!:I!ei! index. ~gcQmes 'mediom.iin.ml fac:lol.
When the LU. ~r:II~rea."iQ:5ri1lild :mnulia!:madliil1lg,

bec!om.e~:mo~e ~eri(YI!$,the gwo'wthrate. is ,d!eWrDined by P~lmitty,pc'. IRS, typical of th~ irnp.IDVI!ld V3rieties,.has a bioo~r opUm~mLAI 0llI.d: a h~gh~r cffi!Cli8IlCY' of dry matm:r pOO;iihuct~Q'~.~. P,~~:Ili a tal] I.eafy trop,i.c..a:Ji variety .. 1lraifi~ driei~ a:t [OS C, which bringsa'b,out ~ig;m:'otls soodlilllg growth. ThLs elU'lygrowlh I3.t:e~ ;;:o.n:Unue:!i.~~ ~ilt>~f staFS. In othe~'WDII'W., [R.:8 hasa
$teiiljdi!y high .~Qwth.rate fr;t)m dte seedUl1IgM~1il: rB han'<lJst. whereas Petia bas II h~Ch growlll. [a:tell·t

Als shown i,liI Tah~le J. m.. much !iiho:r"l:ertharl 8is l:ightexti:nc~iQn

re'la, i!!l'Id has OJ l<!irger ~~'~f b~<l!Je ratio.IJ ~Iso has m.olfC; erect leaves as cx:,plj'es~d bymnea[l .Ieaf

W 1.000

IRS has rernatively beavy grain we~ght. 25.2

openness,. .resu]~illgilllow:e[
efficiel!'lt
V"']l;I~"

,000,

The~

ciI:m:rnctersaru

b('lii~\!\(~d to

.Iml.crcasetl1l! plliowsynthc'l:i.cact:iv]ty of thepopula& tio~an.d!lo mil~rl'taiJaa be~wr bi!.hm~e b.e!tween

ph(l~o~Y!lith~$i$ ru'!:d re~pi:rill:t:iol1.

the e;n:I¥~~ .. ;s fOllliO'wed by !! low ~wth ~·te dl'efn~W~~. Uappellfstha~ t~.e hi~ groWilh :r1l!le: of IRS ,Max~mlilml lrecQ~d!!dI b.AII andl gr~llillili tmel!ghor~t ~u lif~ ~ there:dt of ill. oomlbina:Uon of Viel!d lin~hil! wet ~ii!Son Iflcr~as(ld o;plwmum LA]. and! mmde~ak;l;y .. ady e Fig~re 9 $howc!l the reJ~:ti.Q~'!<~ip'bet\!'~e!JI. [he ~ IlaillXimumoocooo~d l.AJ1 (us~.aUy re:oom~d ;;lX mill.tu,fi,y. It. lis [ogiad to c'Ql1fiider e wh(l~!.!~t~ o:f~e·d:I~f.lgvi&or. muma] ~;ilJdling, a:lIDd lu:ighS[\owlh nower.illg) <JIId gr;ll1l1 y.ie~ilI. i:!'iWe w~t·SieaOO]lCUop. h r~ duritng the Wl!llSllaSOIl wh~n 1he,rveT,Il!g~ !inl:.:r ~i!:te atWa!~:rgoo:w6l ¥Ulge~k! l-efrnilQ'fpl;3ili.t type and malt1i.uity. When a vw:iety Js Il.:'a.fywjth ·d:l'ooPY f.'!di3ti'oll! is ~.ow (ave~Sisilil,s about JOOg·ca1 le;ave£,8:m(lilli!i a, result its optimum. l.AI h;~.s 3. :Iow CJ:Il.'21 d!.lrmg fll,~ .nipe[li:n,g pc~iod) andm,utuil day vaJ1;I¢,. mutuid $h.ullmg!leitS in ~t ~e.I~tilV·dy ear]y :iilimcliilll,El is 3.s:IlJrucnm .pl)oblem. smgcs. and ~he growth irate beeame S iSl:ov.rer:!_tbtelu To oMai]1) ~llfQ~mmt:~oll 011 fherebHonsllip n;i!~. On t'h~ (lithe. ltrmd. W!l.011J avanety has eJect between maximllJ,m, recorded LAW 01,0(1 grain )d~ld, IC:[l;vcs. (a:nldithe, re~iuIDtmg\i'li_ or·Ore Q,ti.m~m UJ. di<lta wer'¢ a:n.a]y?;;ed from various fie Id expeli.!l!lCntll j~: high) 1It1d. ruSr:e]iltiVC~y ~3!dy Iillll,uujng, there is oo.ndllded. al, Hie ~lJist:iW(:¢- the 1966 ,!.tId ] 968 in
~t~'I.Mi~.

In l~~.ease of IRS, .~!l!ir! yie1:(1L~c.rca&!d with c il1c:reasmg tAlI UlM'jl th~ U,[ re-\ili!ih.e,~ai:101)!:t S" Beyo'ndiIJlis vaJuc" &il'ail!l yie:I.d~ccilme: !IJ:moscl COllsta:n,t It appe;us that the I:!:igher LAI did not r,e¥~dt i:Di cl!ecrert!l~d graml1l yic:~d. j ..c., c\xDe$CtiIv¢ LA:! diOJ~ot; .illlVe <I ,tiLetrillm.ent a l: e~rrect~[l.th.c g~a:inyi.c Id

of IRa:.
Or! the ,ather hu,d. the~e Wil!~gOptimi;lm LAII wll:id~ p·l')t!l! ;l:ch[eved .it$[JJ1::lxirmJim gr.a:in. y'il,~I.d. Beyond ilblc l.J~I of 5.:5, thQ gra:in y.ield decreased sha:rpJy. In Qt~~Jt;:l WQ!I'd~~ tJm,¢ grai,[1yireld of p~ ra i:n.c:reased whlh. ~I:l;omas:ifilg LAm Ill.pw ;) .5, then dl~t:rune!l:'lailmutual £rutdiflg .~t m. 1lJIl0i 'gr:a:lin yie;ld d!e>c[~a;scd with rn,creasilill!,I"lLAID,
<I~

_~Al

Fig, 9. R.ell:ltiM5hip b~~weCii!DilX~lrI'illlil'l l~af'M1~il,ind~ lind SHUn :yieMI ()f IRS {l_f1d.P.e;t~, HJ6·6~d! 19ifi~ wet
SE:aWIilis.

23

little chance for mutual shading to become detrimental and the growth rate continues to be high from the seedling stage to maturity. The optimum LAJ of the rice plant varies with growth stage and the amount of solar energy received by the rice population. The reported optimum LAJ's at around the booting stage range from about 6 in Japan, and from 5 to 6 in the Philippines. As shown in Fig. 9 and in equations (5) and (6), the optimum LAJ's are about 6 for Peta and 8 for IRS. The improvement of plant type resulted in increased optimum LAJ. Besides bringing about increased resistance to lodging, this also is an important factor constituting the physiclogical basis for the nitrogen responsiveness of the improved variety. (It is important to bear in mind that the tall Peta plants were tied up to prevent lodging, otherwise different results might have been obtained.) IRS has a distinctly higher optimum L.A! than Japanese varieties under comparable solar radiation. It can be predicted from empirical and theoretical considerations that IR8 may have an even bigger optimum LAJ in the dry season when' climatic conditions are more favorable than in the wet season. It is also striking that a large LAI did not have any noticeable adverse effects onthe grain yield of the wet-season crop. This may have been due partly to the increased optimum LAL Thus, it may be said that with an improved variety, mutual shading is no longer a serious problem in rice growth as it is with leafy indica varieties. As a result, IR8 can yield up to 10,000 kg/ha with increasing nitrogen applications under good rnanagemen t and favera ble clima tic conditions.

Since it appears that improvement in plant type has achieved the goal of developing high-yielding varieties, and that few further improvements can be ex pee ted along this line, this other approach increasing the photosynthetic rate per unit leaf area - might offer a possible breakthrough in breeding for higher yielding varieties. Thus, the purpose of this study was twofold: to verify whether the new high-yielding varieties are associated principally with plant type, and to find varieties with high photosynthetic rates. In our study, the values found for photosynthetic rate were much higher than those reported so far in the literature. The reported values are 10 to 30 mg CO~ I I 00 em 2 /hr. On the 0 ther hand, the values found in the present study ranged from 35 to 60 mg CO2 1100 em? Ihr. These findings led to a reexamination of some of the basic relationships in the photosynthesis of the rice plant.

Some relationships of isolated leaves

in the photosynthesis

Light-photosynthesis curve. As shown in Fig. 10, photosynthetic rate increased with increasing light intensity up to about 50 K Lux. In this example, the maximum photosynthetic rate was 51 mg CO2/1 00 cm2/hT. The shape of the curve remained similar to that reported so far in the literature.
60

~ 50 E
<.>

840 .....
8"
Varietal difference in photosynthetic rate of rice varieties

:if

.5

~30

The concept of plant type has provided a useful guide in the program of breeding for high-yielding varieties. Not much information is available, however, about the photosynthetic rate of rice varieties per unit leaf area. It is possible that the improvement in plant type is associated with increased photosynthetic rate per unit leaf area. In addition, a variety with a high photosynthe tic ra te could be used in the breeding program.

1
[0

/'

..--•

20

30

40 (K Lux)

50

60

Ugh! Intens ily

Fig. 10. Rela ticnship syn the tic fa Ie.

between

light in tensi Iy and photo"

24

Plant Physiology

II~

•
0::

i __ I
3'0' -':: 11--

..; ~o

---

.'

~~,!ii!iilll
F\'i~-,IiJrj""" 1I!iilii-~

--....j..~~----f~~~~----il-----I--~..g

o "b-~'-~~~~~~~-+--4--~~ 'Gil 'i,0 112, :z 4, '6 8 110-1'2:

~
:$Iii'MIJ}limQTIme

O_~----~---~-----d------~I----~I o
2 ~ ~,
!Ii

NlliI:iW,l.efl! ConJefll ,iii leof b!c!d:e

rnl,':

t%J
cO'llll:en'l :lIId,

fig.

II,. :Dii!m~1 ch :lIlse in !lppll!lttii phGlOsyn,U!cUc

af imlillil~d, lea,Yes.

Fig,. ~2., RelatiO.I'ISi)ip 'bcl-n pho,tosynth~:iic .rlll~. -

"iifO!}!!:"

A.ir pow rele, mean COl ,!XJllcentrQ,t'fan i',1 a leaf cll,llmln:r. (mil tlpparem pJl0,t,osyn.r:he,ric mte. A~pa:rent ,photosynUu:tic m,te, utc:realled with imt,c:rca:sing, air no,w m't~ er ifll::~eilsmg carbon d\iiWi.i;dJ~OOI:lC,e:IiI'lmtiOIl1 ~nI s, ~eaf clJ;anWe;r (data! ,are oiThj:t~eCl), BeclJtl~ of dtis, ~ehltIDom;hillp. Ute,
measured phO't:OS)'iR~U~ ~a:t~
'iV1l,S .~

~g

......

s
.!l
gi'

40

.... :;10 ,aCllli:Jtst300 ppm ()if the mean c-awcm diOxide eon,eentrlllCion mtlte' I,ear diamb~r. u Tltw'! of mmpliirg;. Whf;1iI mnp]iLn:g Itodk pl,IIQB E several hours ,afterr sunrise. thep:laJlts oftel:lsholNe,d iII!' :20 I SiglllS of 'Wiltiing., They vbj,b~yreooy,erd INn,en ,ti! placed in a diadk,room 'Of !.!Il'IdeJshlldec for SO:Jille ~' hours but tlJ..eir I?ho'to~yn'theti'c ~ates r,ema med I'ow. 'E. ~ (I oon~razy ~() what 'WlI_S, expli:e,ted. F.ipre .~~ 5how~ tliI..e dl;IlIi'ti!a1. chal!l:ge:in aiP'parent pholloiSY1rl~heti<e .r.u~:. 1he: p,llmts &il:mph:d. e3i111y .iln. f. I the moml:nl8, or I:at:e'in ~hii ,~venmg ,showed 00]]~i.$?tentlyrnglil. !!'ah~e$· ,d1these sanlpled iii lllli.d-{hry M
,

s'Ulndarliiized

a:
...

r:r

y ,
•

'

. / '.
I

• V

.' V /

ill' £1;

!!;!I

I
1

mQw(ld ~ow val,ues. TlW: variation inphotosynthetlCr3.'lC is pfobo:!bly relat~dI to the w,ater supply of 'the Iphllll'l. This :illldicatesthat: sa~pling5houM be, done early in '!he mornirll,g 00, Qibtam the max:imum lIctiv,ity of ,ho~'lhi!::iis of ~SO:lated h:a,ve$~ hi ,atldlilion,. ~ady mo:mIDng sMi'lpUng, ,gi¥es m.o~, ,ollOsistielil~ redts, thaa:! .ntid.-day sampling. .tm. this e__;(pe.riir.m:l:l1,~:, the pliin:tllwe~e ,samp~ed ,ar~,
ilIiiound 6 a.m,

Fig> U., Relali,Dn541ip ~tWtii:"

,iflhl:I<tO$YJiI,(Il:i:~il: Irate.

clifow,phYI~ ~rjI!'Ulen'l:(lnil

Iffethad of filQUn.lIJ!g /eqf ~,lef ilil Q ,I'ea[ Chal:1ib-el'. A leill bt(!.die 'was ~[llIU'l1l'ted. from the: :shll;ltffil II,~,(I ,pla.ccd. m ;iiI snlSiLl hlle. Tiiu:iis ,ope~aJUo.n.

was, ilccomp]ish,edilltl, walte~ roiillow

tHlltliOU,S

fior ~ con-

now

Clf wa,te~

mfue

trn~ues. The small

Plonl.'lrysiology

25

18

~
!ir

!
800 <:

16

If
14

'"
E

• • • •

'" s

;;

12

.J<! 1!
1; >. 0 <:

"'"'"
E
1&0

,0

00

..
zoo

00

- . - .-0

..

• • o.
0

.,
li:

1O

" Z

t .0

r.
Nit""J!"l

Con,,,,, 11'9 N 15'1

220

240 em)

200

Fig. 15. Relationship between nitrogen photosynthetic rate of 50 rice varieties.

6

content

and

(I
30- 35 3~-<l0 <10-45 <15-50 50- 55 55-60 60-65

ff
-3
Q

860 ~
",.0

00

Photosynthetic Rote ot 60 KLux (mgC02/100sq cm/hrl

s
Ii!

Fig. 14. Distribution eties at 60 KLUX.

of photosynthetic

rate of rice vari-

..,

,l!40

·• • •·••....• • •
0

. ,.
0

••

·0

o ••

1.

••

I.
tube containing the leaf blade was placed in a leaf chamber. Nitrogen content, chlorophyll content, and photosynthetic rate. As shown in Figs. 12 and 13, the higher the nitrogen or chlorophyll content, the higher the photosynthetic rate. A high nitrogen content was accompanied by a high chlorophyll content.
Varietal differences

~
0.3 0.4 RoIoti"" 0.5 c!>loropt.ylJ

•
0.6' 0.7
0.8

Content (una oreo !>mis)

Fig. 16. Relationship between chlorophyll content and photosynthetic rate of 50 rice varieties.

,.!J

Photosynthetic rate of 50 varieties. Fifty rice varieties from the world collection of the Institute were grown with 0 and 100 kg/ha N on the Institute experimental field in the 1967 dry season. The photosynthetic rate was measured about 30 days after transplanting. The photosynthetic rate of the varieties ranged from 34.5 to 62.1 with an average of 46.6 mg CO2 II 00 cm2/hr at 60 K Lux (Fig. 14 and Table

. ••
'5 -j!

-.

..
0

•
0

.0

·0

.- .
•
0

••

••

I"'"

1.~~·~Q--------~--------4~P~------~'~.3--------~~O
leoI ~ ("'9/"1
ern)

Fig. 17. Relationship between leaf thickness and photosynthetic rate of 50 varieties.

26

Plant Physiology

mg COi; I] 001 C.ntl:! jllJl'·. U. is obYi.oru:s fmiIU these findin![lS that~riela1 wife.ren,aes exJ.!s;L in ll1i.e ,pllio'tQsynthetilC ~t~ of f:ice:va~[e:U~s. Nirrogtt!' content. ,chJorophylJ ,G10~t~.I'U, teaf thr£kn,e£r~ anti pMtQ£;}Ituhenl; m:te. Asalrcady ~hoW'ti i!!IFi~~ 12~d 13 &ere :.l:re ,clQ,s:ereJat~on~h:~p$ b~~.ween[(lhot~ynth~d~nne!l1lid .n.itwQge['! o.r .dilJQ.fO\phYIW ,oQn~el1l~ ith:in the sanm:var,iety,.l:e<llf w

4). M.o$t V;'Iruu~~felJlm the'

rM&l:l

of f;fo:m.~O ~o S5

Fig~re~, ~5, 16 and ] 7 mO'"w th~I'~I:atiDnsh~p~ bet~~n photQ~Y.!lthe~tc~~af:ld. .~:l.trOg,e:1il Im.d.

c.hk!.ooplruyU ,eonb~J:IiIsand IDeaflhjclmess .. 'fhre 'm, lo~ (lorrf:eIO'!tiIOOiiS betw~e~ !phQto~y~t.bet!ic ~a!t\i!l Md njt:!"QgeWi. 'OQlllre[:!t. dtIDoil'o.phyU c::<m,~~',nit,.Of ~ear
Ultidc:mess;,Alorw plmtosynmetic rOlire is ~oc~~t'e:d

tbick!'l.es~ is C(ijl.!'!s~der;fld .~ ene of the f~ct:Qf$;tlli!~ :afFeot th~absQrpti.()mI ofliight ~y 11 leaf.

wit:h; .Law ni,trogell c!(ffl.~:!th Low dIilQ:I'ophYJru·ool1!' l,e~~.a[l(l .. Iow leaf ti.i:CkmeJs"s, VMILI,e:s" A h~Sb photQF synthetic rateJs a~~oci:atedMIt JUgh~jltrog~ COlnent,. high cll.IQ!!'op,byJID L'lQl:'!bmt. IUtd high leaf thndk:ll(l;s:!l val.u,esi.Howl!,w:r" theee is 3, LaJ:ie vMiation

Photosl~mthetic r..u,e
Itxpt;. .IiIO. Val!'li\~w CmJl!1llil'Y or

~COl I:T'iIg/100cm::l/hr)

N itro~rn Chiaro·

baf

Hl' K Il~x

160

iL.1lI)ii,

,eontel"l!'[
hng~lcm:l:}

phr'(l ~~hr

1~~.jIltion

con~~n~ (mg1cm,2: ~ ~.~

ckness

'1

rCalaro U

H4

AU~lraUa
,Qeylol"l O~yh::);Ii'l ~Vlol"l 'Colornbiil 'Colarnbua ~rg':{pt

Plilkkalli Paohchai I?'erbumail p,~163540

14.0 16:.4 '18,6 17.0

3Et2 44.9' 43.6

51',8

177
219 2'25 20'1 230

0.5S!
0.53 10,55 0.,.73 0.71 0.,.77
0,52

3.90 4:.l:i:5

4.3!5;
4:9t5

:22,3:

()I :Bl08
Gamb:r~h Se:~la
Araby NaMal

20.9
20",4

:22.7
Ut,S 19'.2

T141
GEB,24
Biiil51il1ilaiti

Egyp'[ Egypt India

Ind~a
IIl(;H!!

2CM;1
2U!
22,2 2~1.6 18,3 24.3 29.6 23.,3

44,1 5Ct4 49.3 43)1, 44.'0 62.1

4 ..301
4,.50

209
164

4.40

0.46
0.,57 OlJj2 !174

3,516,

::210
19~, 172:

0,,130 O)~H

JI'ii(loa349' ADT.27 Bjll

India

47A3 52.7 0.:9

47"0' 55.6 47.5

ass

4,;40

2:2:51 'U!91
216; 201

;,;'BO 4J35 4.40

azo
4.1 0 4Jl5
3.7'0

Iru:Ha
India

teo
UM
1'77
'1912'

1159
O.,iEiO

Sig!lld~s IPll'~:a 8!e!il9awaFil

Indone'S!'"
Irl!1:ig:r1Ill*lij,a

(l.57
0:,58

46Jl
45.0 47.0 44.5 42.3

me

I ndone-si'O! IflRI

U!r:6
'19,7

200: 179
23:4 226i
1

~R5 3558

IRAI
hOlly' hairy'

201,9

0,5'1 0,;62 0,60

0,56

4;015,
3JIO 3,65;

P Ros:s;i
Str~pe136
Yab~mi 1!!5, Flli!! isak:a 5 DlIiik,oku

Ul3
18.,.3

haly
haly

20..4

":1

l.50
4.45 4.00
3.815

44Al
43.7
46.,0

193:
190
201

Japan
J~p:an:
,JIi1l!ll:a!"!1

20J5

UU'
13.7 22.6

42.6 39.9

HO'VOik'ig!1

3~,5
49.2

172
219

lIankai: R orta1n

M!!~'!!'Y:~i!!

0.66 la.53: 0.73 OJH rQ,66 Q.5'7 0.60

4.1];54.110
:3.415

3.7:5

27

Table 4. continued

Expt. no.

Photosynthetic rate (C02 mg/l OOcm2/hr) Variety Country or location Nigeria Nigeria Nigeria Philippines Philippines Philippines Surinam Surinam Surinam Surinam Taiwan Taiwan Taiwan Taiwan Thailand Thailand Thailand Thailand U.S.A. U.S.A. U.S.A. U.S.A. 10 KLux 60 KLux

Nitrogen Chiorophyll content (mq/cm") conteru.'" 191 203 205 220 224 194 181 181 185 212 202 214 220 200 208 227 181 195 250 196 242 223 0.61 0.60 0.57 0.51 0.38 0.50 0.64 0.47 0.56 0.51 0.67 0.62 0.55 0.61 0.50 0.60 0.45 0.53 0.50 0.53 0.49 0.56

Leaf th ickness (mg/cm2) 3.90 4.15 4.10 4.25 4.30 3.85 3.45 3.70 3.55 4.20 3.95 4.10 4.15 4.35 4.00 4.50 3.80 3.75 4.80 4.05 4.70 4.60

29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50

Donabaru Suwedee Dikwee BPI 76(1) Milfor (62) Wagwag 81 B25 SML·Apura SML·Kapuri SM L·Temerin
Dee-qeo-woo-gen

Taichung (Native) 1 Tainan 3 Chianung 242 Leb Mue Nhang 111 Leuang Yai 36 Nang Mai Siam 29 CP 231 Belle Patna Nato Blue Bonnet Min. Max. Mean

24.4 22.7 20.9 21.2 18.6 17.5 18.5 18.8 20.2 19.8 22.6 21.8 11.9 16.8 21.6 17.7 18.2 19.6 20.8 16.5 19.7 16.7 11.9 29.6 19.7

52.7 51.1 49.0 47.1 44.5 42.1 44.9 45.4 40.3 47.3 53.0 45.2 35.3 43.8 57.8 47.2 46.2 45.6 55.0 40.2 54.3 46.2 34.5 62.1 46.6

* Mean values of

a and

100 kg/ha N. its biochemical activity. To understand the factors responsible for varietal differences in photosynthetic rate, a study is underway on the photosynthetic rate of chloroplasts isolated from different rice varieties. Analysis of Yield Components of Improved Varieties In the 1967 wet season and 1968 dry season, various field experiments were conducted to find out how spacing and nitrogen application affect

* * Relative values.

in photosynthetic rate within the same range of these factors. A comparison of these relationships within a variety and between varieties suggests that there are other factors which control the photosynthetic rate of the rice plant. The photosynthetic process can be considered as consisting of two parts, namely, the diffusion of carbon dioxide from the atmosphere to the chloroplast, and the efficiency of the chloroplast in the assimilation of carbon dioxide. The first process may be mainly concerned with the anatomical structure of the plant and the second one may be largely dependent on the micro-structure of the chloroplast and

28

Plant Physiology

tnlegwai:nyje~d.~d yi~]dco:_m~Qln~~~ of imnp!fo~ed

~Q"W-:l!l1!d

'j,g 1F""~""F'~"---r;I~~~~'"T"-----'r----.------'

.litifll'htiIDlflrlQg,Y!lrieti'l'!'$,~rno'w Is an 3i1lalYlitLS tbe graffin yic:.lds fr()m,~se !experi:menls: :il'lkmu of yre:Jdcoml:ul'ne:n:t5:.

or

.~ __~ __~1~_~8~_~_' __
I "

!l:igl

~O __

~D __
:.

__

~,i;stcdm T1iII!Il~, s. ifi,w!i'BSie:tics~re used fin a!I!I U~ e;(re'.!'IDm.-e.!I!~,wath I~Wg, ,of t.hem,lRS I(VS) ot._llii~ .1 R.~S44S~1"\3-3 (V2). b!!!imguS>!!Jdj ~'Xillm:p,Jes as .!rw. tih!:l'_'~,aJym ,of yielJd! c@mpollle;n,is. Th~\I',Jl1i.ela], ,ch<!"'.!ilc~!dstics.Qrtilile~l,\'l.!rie'tie5 We~.e d~miib~dI 1m'! l!:he~.961A~I:IThIJlSil Report
Palili~le, !liIumber par :squam
!III!lte~ WIpillllyieid

1l!JLl:dia:ta we~'(lollected

from the ,expe~~ts;

~ 1--+--Ic--1--

~~:it! Wjjl

~D~."';'.'.~

iMiiiP:¥l

.-t---;

!~O!!---",!iOO~ ...-__'" ~~)o"""'--'::lilO'!'.""'. --,_~.c-. -~'500~.'_"-_..l!. "....--."l~~,

II

~:(ilI:~I~m

IG!l'ili:~ yi.eld wti~~ooely

~daJu:d to ~.:ic:le £In;uJIlibe,r pers:q:1:.!a:re meter upt(l ;!ib{)lUJt 350 to' 400 .~f sq .!lru; beyondtb:is. grain yi-eMwas liIottwe!~yre]a~efJ H'I pafilJi~le f.lll!lmber (.f.i;S. 18). The dry.:sealiol'l c~o'p p.rodilJloed mate pan~d1e~ U!m~e wet-~O!SQn C~(:Ip am,d the ~fle[!(I!a.~d numibc~ @f pank:Jcswas !!.:s.sOCiliil~,eI~ w~thi!lJlc[eilSCildpai:n, yicJd,. l'he diI\~ab seeded crop ~II'OdU.c~I~i11 much ,mgew lIumbew of pi3:!Thides th!l!f! the t~omspl<mb~dcrop but this iill!. enase did not ~:rm,B. a!bmu~;e ~~cfl!tascd gram yieMls. 'fro!!: was beCilIL!SeWe m~reOl:le .m th:e
:~umbe:fofp~~cles w~ !l.ccQ!!l:l[Ililni~d by :a d.cC:!I'caSfl

FIg. ~8. Rela,li.o!!1$bip Ii;i.t:;hyt~iI ~Ulicl'e Dumoor'pel m~~ef !lfi.(lurun 1Iield.

$iq;!i.!~

in tih,e: n1um.ber of grai RS ~:r p<l!lli:~le (fig. 19)1

Paliliicle n!ll mkr
!Il~llfIbef n~!e

'of

persq,QO'I'R! nM!!W q ~illil pet:'. lpalliielll!!

,,0

J1.1!!I.n1iber otgliain!i

~,!:!llltedto

pe~ JP_:ide: i~ m,ve~el.y wU~~lI:Imb~'.f@€pwicl.c:\i:p!![ SqJllISite

5Q~~"'-~-+-+1 .. ,--~~~.,-~.~~,,-~,~=_~, _.~ ~.~~~~~~. .. ~ ~

~I-I
°O~·.--~I~~.
~j~t'lQm

Irfl1!~J9. Rela,tjilll~pbeLWem!
m~til'Ji' und. Inulllli!Jer

~rgr:fuI~.pei'

p~l1I:icl~ I1iUliIlbm ~ pD~I.e.

~.U~

Nitrogen
IEi'i'pt.

:Spaci~:g I,em,) Of
Siler! ingl r,aU!,
1~lk9Iha~

no.
11

Y'I'!~J

$t!a$On W'!;!! I[),~¥

rllill1l~ingl me1:hod
1

!'Ipp~i(ia~T:on Oll!l/ha)

~~US:7 UI68 1961:1 1008 19~3'l3

Tt,Blii'I!$,lIl:pruITii'IIQI O•. UID' Tr.,m~pllil:n;~ITngl 1'.r:af1l,~pl!lln~UI'l!i1

l'r.af'l~pl<l!l'lIling, 0,. '1IO@r

11Ii! " 2'(1:l:, 302: ,.40~ ,. 'SO~ (

2 3 ·4
I) (I

1Q<, 20~. 30~"40itl, 50~

20~ 2G1 10~. :0~ 2

Wet.
Dry Dry Dry

O. JiO!. 60 .•'90, 12iQ

l'r.a~~pla!'llll~n!:l 0, 301 60. '9@\ 1::;lQ

0.1' 00, 1'50,200,,25D

130'

1968

Dlrec:~'!lE!edIi!'!~

50,H10I200

and this increase resulted in increased grain yield. This linear relationship indicates that (I) the total number of grairts per square meter determined grain yield and (2) two other yield components, filled grain percentage and grain weight, did not affect grain yield. Filled grain percentage and 1.000-grain weight As shown in Fig. 21, the percentage of filled grains and the 1,000-grain weight remained fairly constant regardless of the total number of grains per square meter. Therefore, roughly speaking, in the dry and wet seasons, the relationship between grain yield and the yield components can be reduced from:
Grain yield ~ panicle number/m 2 x grain number/panicle x filled grain % x grain weight to: Grain yield - total number of grains/m2 x K,

o Dry season • Wet season x Direct - seedl nil

10

20 Totol Groin Number / sq m

Fig. 20. Relationship between total number of grains per square meter and grain yield.

meter, i.e., the more panicles per square meter, the fewer grains per panicle. For example, the wetseason crop tended to produce a larger number of grains per panicle than did the dry-season crop which had a larger number of panicles per unit area of land. The direct-seeded crop produced a much smaller number of grains per panicle than did the transplan ted crop, which produced fewer panicles per square meter.
Total number of grains per square meter

where K corresponds to the gradient in Fig. 20. When grain yield is relatively low, panicle number per square meter can be a good measure of growth performance, but when the yield increases, the total number of grains per square meter is more reliable for comparing growth performance under different cultural practices. When trans-planting and direct seeding are compared, total number of grains per square meter is the adequate

100

vs grain yield Since the number of panicles per square meter and the number of grains per panicle tend to compensate for each other, neither of them can be an adequate measure of grain yield under diverse cultural conditions. For instance, close spacing can bring about more panicles per square meter but this increase would be accompanied by a decrease in the number of grains per panicle. However, the product of the panicle number per square meter and the grain number per panicle, i.e., the total number of grains per square meter, is closely related to grain yield. As shown in Fig. 20 there is a linear relationship between total number of grains per square meter and grain yield. This linear relationship is independent of season and method of planting. The dry-season crop produced more grains per square meter than the wet-season crop,

q5'
~ ·0
c;

80

••

J~~
"

&0

<Q ~

<i;

~ 60 I----- 0 D.-y season ;;:: • WeI season • Direct- seeding

..,

I
fliP'

& 6
Q
o

"'~
20

,.,0

"

no

v~

20
0

10

30

40

50 XI03

Total Number of Grains / sq m

Fig. 21. Relationship between total number of grains per square meter and filled grain percentage and grain weight.

30

Plant Physiology

T!iblc {i. The modell G'iillin

)fiel\!! COIII,pmw!ll!lI!

or lii'ii!iispltinledl
lOry

IIRK

yie'~d
6,l63 250 1100

compclntlllt

~.

!
>'-

,~t--~--+---tr--c.7T~~~--!r---"""

,Iii ~ I---I---::r-Jto-ly'::-t--+---ii----;

Grain vi,eid (k~lha'l' ,Pa~iClr,eQ11l:1 m.ber.l$q m IGir<:liriil fUJliiilbe#'p'an icle 1"01011 grainl numllXl~{$q m
Fi'III~191:aln (%1

9,244
1

25',000

~.OOO gra,il'ii weiiglrl:t (gil

as 29

UlO 31,.500 135

29'

315

°O~·-~~-~~~---M=L_~-.=~~. ~bl'lt ln faril1Ilr;s. fieid.$. --~~~.·--~~=_X~~.

lotal Gi'ai~ ~
I
$iQ "',

cOilltmlll:in:g the tiillering p'e:rfo'~i1.oe, of the rice

:F:iJ!.n. :Rcl~lign$hip

I':!;ctwt~ 'k1lall'liimber of grai!il$ ~ lQlI~~ meter !lind .",,:lilll )'iel~ (If two· varr:ie<lics..

meawre

t:or ,evm\l~U"lg growth

pe:rsqwre:

pan~cle: number

m'CEcr

pe:rfQ,fi11<i1:1Ue;.ll> i win give less

iR~or:maUon.
rrlKuJel yiield eompolilellb

TheR are ·col'ltr.adh~:·tiflrg·I)plnio~$ as to' wbeU'l:e:r (If !l1iuogen ill the plant IU the !!'~t.e of nitrogen ~phtkeper day is reila~ed ~o U1e ~.i:I'le:riiIll:S, pew'Jicu:mance, oftlliiJ:!"iee Fur;tlltl~~ mQif~" nICe Va.ri~tjIl5 haY'!) l,ul!:elvJ/~~~a] .eli [€el'Ulces m tm:e~!, abnit;y. the natlY:re of' whiicli. hm's not: been ullde:tS,tood wen. To iiletl:c:r wuwrstandl these;

·the: ·concentratio.n.

,mant

IiIlSiUer.;an tlixpcnment Wl!!S condYided of 'b'3nspfanted

IIR8:

ro SltUdy(

1)

Table 6 gives a :roup ~deaJof :the yieM oo:m:!lOllen1:s (lIf t~a:nsp~an~d ]ItS in d~i) w.!)~~Emd ruy-:s)e'asom!.
·OI:Ops.
Co.lIilpadfilgl ~!Ji'Ef;er!'l1It..,ari.~iBs

'[he lI'e~aitiolilshiiptll:tweeml,Wt;fo.gen liIutrition and UIIDertng perform811.ce:,. and. (2) varietal diffeJ,enees :in tllilerir118 peirfoiltlil3iJloe•. 'The ri~' :plan·t WSiS grown ;a'tillir~e[~JlJi~ niUQgen. li~]S usiJlli!I the :s.t;mda:rd ~1ler .wltureleclm~q~e. .!l!!'Iid 'the ne~essmy da."la were colllect.ed e~ wet:k from 2 weeks .after
U;ii'lS1)dMJl:ing. Dlange ,jllli tin. nl:lmb(lr III; dfected by N l'iill'emsin e<u Ituli'e sOlution As mown in T,tb~e 7 the iniluatioo. uf tillerin,g was accelerated by incr,easecli C,Qucen.ra'UOfi of n1ttog:cn~a1:so, Ute Iirlgher the nitrogen S'upp:ly, U!.e' Warger the tiller numliloil' ail: ;any time. This, ,deady mows ~ilIt w'lII'o,&en sup~Jy oontw:is the liBeriAg fi'!:t~ of tllc rioe .phmt lillP£ess ·ctlIDef fa.ctors such as ~p3icm:g~dl bcco:llne brl~m3.

Since gram, yieru~ iiS ;'Ippmx.iIiI'liat.Q·ly the p!:'oduc[ of

tQtl1i11ll1ll.m1D'l!:f O;f'S~ailil:S:per ~~Ii!arc meter :uld gmfn. weight. and grnii~ wel:ght iis lnllillly:! V3luietall
Cha!l'BCtilif

and

is. .not

cllt:m:8.ed much
Ci)Jld!iUons.

~y

cu,1rurall

practices

under

D!!UiUm

it (grain

weighO beeomes aill. ~mp'ilIiWit factor ~1iI comparing d'iffe:relllt varieties. Thb .~!.lla.·UO:I:IS:hip is sBlOWlil ;ilil Fig...22 .. Tho ·Y.Blriety 2 ~.~ O! ima~Je'f .~,a:iml weight V thaJ'i 'IS. CCiIl:s;eque:n'IJy,. V2 :ne~eds mere .gr.aJ1!lIiS ·to obtlIin t)1:' $ilm..e .(lTaii~ yi:~dd. ,as VS. There Js a t(!ndemllCY iOf alj13liety of ~anle:r Bli'.rui:n. .e.illl,t. to w ,lodu:cea I.a:rger number of grai:liis pe~ squlU'e
meter.

Jj.~.

'lotlll

INlliipUlkie VI

NI CQn~Ii'i;t!rt!tiQIili

Effeets of Niitrogln INlYltritio:n 'onl

Tiilll'sf'hil9 ,'erformance
1

In 1967. U10 co.l1I;celd ·of ire~alive Ullerilllg r.ate (RTR) was mtrodl~Qe·d 'to :s;tudy the effeds 'of :w:lnngen. :pbospho.rus, ilfIdpo~lIm $lWPP~Y 0111 the· ti1l1.crjngperformarlt:e or d~e ri:~e piim.t Of ~e.se n~~:rients. !il.itrogen jlS the mos.~.d,omrunant iml

At tliI.e early ,gliQwth stages of therioe pll1il:!t" l:illle:rin:l'l.i~, f.lOli lI1!il,ated.to' tkemtrogeIlC(lnCent~,a~ 'lion in the p~anil:(r.~bile 8 3II1Id Fig. 23). The d!!ita millk:ate tha~ &he:rice pLant started 'tillierin.g, wihen tht: tot:a.lim.tmg.e.n uptru became :moJle 'than lO mg per .pL_t Or th~ dO' we;i.8b~ ltiIe·ca;memo.re lham 300 mgpe.~ plant.. :In othefW(lWS" ·tlle:si1Je,of the :Jm'iam def de'tenn:imuid (he ;im.iiHa!tion of tillcw.~;fl. When the mmtme~ J$. mo, srnal'l., the: w:rp:lus, ,cOiilibohydil'aibli it plI'cu:liuce5, is cQ!:!$u!l1ied;mamly for

:u

Table 7. Effect.sof nitrogen concentration in culture on tiller number of IR8. N level (ppm) Weeks after transplanting

2 1 1.2 1.2 2.5 2.4

3 1 1.7 2.5 4.2 4.8

4 2.2 4.1 3.7 10.9 11.6

5
No. of tillers

6 3.5 8.8 12.1 24.6 27.9

7 3.5 11.5 17.9 35.3 37.2

8
3.5 13.3 22.6 42.7 44.7

9 3.7 13.9 30.2 47.0 51.6

10 3.7 12.8 31.8 48.6 56.6

5 25 50 100

3.3 6.4 6.7 16.9 19.2

Table 8. Total nitrogen uptake (mg/plant), nitrogen content of leaf blade, dry weight vs tiller number.

Weeks after transplanting N level (ppm) Item N% in L.B." N mg/plant Dry weight (g) Tiller no.

0 3.20 0.43 0.02 1 3.20 0.43 0.02 1 3.20 0.43 0.02

t

2 4.81 4.44 0.13

1

3 3.67 5.85 0.19 4.36 12.0 0.33 1.7 4.42 12.7 0.37 2.5 4.09 18.9 0.70 4.2 4.57 23.7 0.62 4.8

4 3.18 21.0 0.85 2.2 4.25 0.90 4.1 4.80 36.6 0.90 3.7 4.46 132 3.51 10.9 4.76 158 3.87 11.6

5 3.63 29.0 1.03 3.3 3.48 107 3.63 6.4 4.50 113 3.94 6.7 4.45 396 10.8 16.9 4.32 353 10.8 19.2

N% in L.B. N mg/plant Dry weight Tiller no. N% in L.B. N mg/plant Dry weight Tiller no. N% in l.B. N mg/plant Dry weight Tiller no. N% in L.B. N mg/plant Dry weight Tiller no .

4.81 5.39 0.15 1.2 4.91 6.16 0.16 1.2 5.13 11.1 0.31 2.5 4.97 10.5 0.27 2.4

25

50

3.20 0.43 0.02 3.20 0.43 0.02

100

.. leaf blade

32

Plant Physiology

Ihcrcr.JJie, RTf{ is the rne:l!fQti)1ierll"!\8 [:Ol:~.epefwll~r. l

--100,

i
~ 110
IFI ~~~

..

1,,5 1---

00--<1

!l0

__
V

'.-,_' ..5 '

1;"

, 2~

Ij;:

:~4
Oll' 3'

M~e:R R'f'R was ploued llga:ins~: tb~ 11Ii:tmge[! ,conumt of I~e: leaf rb~3de. th~ R'fR tellJl,d,eif!lOI be sm,aJi!ef ,a!t(ffia,e: sallil~ ~it~loFI'lCiOJu:ent wile-I:! the ti~lern~lii.mb~f ~5 large. This lllI:I~~l~ tJi1;a:t 'when fhe lilin~!!' l~lilmbeJ:' iIllCJeill~es,mu,Ul.ai1hading wHhilll, $ t~e h.iUretl'!i!:d5 tiI]e~pro(llJicU(lm!,. 1',o~es:l t!a~1 ,abcr¥e pOS8i~il;ity, R'I'R 'Was p']ot~d ;mpinst tililIDer IlILuubcT. Figur:i:l 24, Lrad.icales tha,t: Rl'R is Itiepe,l1Idel'll on the totarn tillme:r 11!lJL[t!]j'er ,.resent; tne! hLrg.er []~~ tiU!l~ I!i~mb~r.!he ~mal~er t the R.TR.. This ~,~~t~g!'ls.hi!pind!kil~es UtM. m~~uI!'!1 ~hlldili!'lg w]dlin th~ h:iU js ;;mi!lT!lpo;rl:an.~ ,r!lC~!O.r 1l1fl!llCtiQg,Ilirul~rir18r'!:le,. U has been cs:lf:lbJishe:(I tiJ!t<tt light i~:rcqu,l:md fQf~ilI~e;[devemoplfllent
1i1n.!s, R.'fR ls detcrmilu:d

oonlont but It~$Q~y tJil.e: present

~!:)t;d IlUmbell"

!lo,l Q!llliy by :I1li:tKlg~[1 of li]I~U'S,

70

Ni~ge;till 'COm'telilllt ,of I~ellfWilde ~m.d Sitr~w !il'sR'l'lR The ;prece~i"!l, s~liIdies hav.emadle:clcllf that fI) Rl'R isi~dc~,fld0mtofl:h~!tit~og.e[! ,oonl~Je!nf of the plant until Lhetotal
mOire

nil!l'ogC:fI t1p takc ~.crei!:'le$ to l~hrl1f11~10 .m.g, petp]~t.,ood 1(2) R'l"R ~ends to
i

~ec:Q,!fIl!c MtI!l!.1]e~ when the lOlail

tm~f

j1!l,!mb~~ ~~~

.brge. The&(lJ fa:t::~(I~; were cOIl1~d!e!l\e~wbc:rl the ['~I,iltiiDn~hip bMwecll R1'R. :arrud the !'!~Jt;J'ogen CQJ1r ~\l:It of th~ p!i~~ 'W1l!$O,tlght As shOiwnirilIF~g;>. 25 S !liQd l.;6.I:iI'Ie~r roe:laHonmips e'X.ist betWCilllRtR .md Ilitwie:n (;Olillielil~ oftJile :1~3f bl~de Qr I;e:av?s. the np,lm ~liIdIDcmlt:c that OtL'):l1JitrGg.en OOI~~,rnF
HaJJIi!l1,tl~e plant co:!'!t~QI;s ti:l~er,i:ng :rnlie!liftc~ th~ fig. 21. RC~~itiOJ!~td,p ~tMl~nlm~f l!Il!liI]kr, r1itto<g~n ,~{lJ'I~enlof Teat II:!i[!;]d~' Md. l<l~31J: rliU'ogen, I:iPlt:ll.1ill" l

<4·!i,%1I!1 !l""l,~t'o~

amid ~s not ~iuffk:ielJl forlhe lP~gdl!.!>clffion new tmers. HiO'WIeVCfwh~'~ th~main of , ~:illill!,rbooomes :l,'DQIlI,ll3 OO mg. the w~~iu!1> UriDOr hydfl!~e C<!D ,mppoR 110~ Qlitly Hs own ,gr;ow th but ailso tllIif)J pmd,'IilI~lioilll,olr tiIDle~s~
~~~,own ,g;!''Owth
I

• !;II
)I

2~!%N

,H:!%'"

liner iIIlllmm:berllsRTH RJ:)clati~ li!I!Jler,illg [,!I~C ~ defitrn,c,()j ;as:

RTR'''---:;''"~~

d.I11 ,d!

I, N

O"~~~~~--L-------~~--~

10

!O

40,

~o

Tillar INumb~

wIit~rg N Ic;fe~s to uIIIlm,b!!!'[of ~:iI1~.eFSU~, to' I~ime. t

FIg. 24.

lirr~t:s eJIf t~'~'llJI WIll<Jn~mb~t Or!ItIR.

I,O.---~~~~~~~~------~--'

Y·-0.3901 r: 0.8832

+0.1988

• •

and 0 ppm N culture solutions, respectively. In both treatments the tiller number increased when the nitrogen content of the leaf blade was over 2 percent (Fig. 27). Tillering stopped when the nitrogen content of the leaf blade decreased to below 2 percent. The experiment demonstrated that the nitrogen content of the plant is an internal factor which is related to tiller production. Varietal differences It is well known that there are varietal differences in the tillering ability of the rice plant, as shown in the example in Fig. 28. To study such differences, a high-tillering variety, IR8, and a low-tillering selection,

2 3 Nitrogen Conteol 01 Leaf Blode (%1

Fig. 25. Relationship between RTR and nitrogen content of leaf blade.
',0,------------------,

~5 e.... -8 c
~

CD 4

'0
Y =-0.2718
r:

+ 0.2046)(

c:
0

~2 c:

0.9108

U c:

_g

80

• •
1:
o
2 Nitrogen Cornelliof Leaves (%1 6
L..

40

30

:J z 20

E
L..

I-50

ppm ppm

Fig. 26. Relationship between RTR and nitrogen content of leaves.

i=
10

050-0

plant has reached a certain growth stage. Figure 25 shows that RTR becomes zero when the nitrogen content of the leaf blade is about 2 percent. Another experiment was conducted to test further if the nitrogen content of the leaf blade is related to tiller production. The plants were grown in I ppm and 50 ppm N culture solutions for certain periods and then transferred to 50 ppm

10

12

Weeks after Transplanting

Fig. 27. Effects of supply and stoppage of nitrogen on tillering,

34

Plant Physiology

lR 15445-1-3-3. weilie g;fO>m m c'U~tlJlru sdJllti!anws wilh djfflf,[~n~, ni~ro~t:I~IU:p'P:ly. A£ shown. ~[iJ F~g. 29. with.in thesam.c:va:Ii¢'~y, RT:R. ~ 3. Im!(!;lt!,' fl1~,c:tio!il!~fth!:l n~trogen cOWlleiliU of the leaf blade,. Wh~rn,low- $Id nigh-till.erlngvOlirieItie~ OIJI1ecompared. ,l:uJwevcr, :n:i:lil'o,gel:lc(mte~t is
n:o~ thi~ Q!l~Y important r~ctOir Call.!sIDJli8, diffef'flnCeS mtiJillerimig ~cr,fbr.t:Wa:n:ce. n.e fact thIn. !'Itm~ sa:rn,e

m.tr~g{ll~ {l:O,IUe:!!!t:.l;h,e hjg~ ~ile:r:in,G,"'.<lindy had 11, htig,h RTR, ruggesls [ha~ odlJ~r f~~tQ~ $ych~s hOmiQfi:e~CQ;Il!t~QI !:le: tilJerilij: habH, ,of ,il!Jff.ere:n1
vmeti~£., SallinH;y pli\obl~eoo [iii rilce lil!lilrti'ii~iCQflI

tn l!. f!reviousUil!!diy of l!he effect 'of high COnteR" llrlli,tioll1liS of salt eariee ,~owth" somUiflill,ulitl.or:i(le WaS usedJ<)I$ tl!_,e ~es;t $;liJ:t (1967 AnIDJ,aiIR.eport). Undcf nal~~aI. oofidlitial1:s:,iI!Iite trn<ts o.f salt diffoi! f~~m one pl<!.CeloMother m, ~aill·iil!fiededaTe;a;s,.It is,Uteircfo!l's..oe~essaJ')' tokRo.w ~e ~f'~c'ts of d:iffe:rcllt i!:ll[ld!l OfWiiW Of mli/dJ!g,res o:f t~cm QIl riee gm'OWUl. O!lth~ (lIther h~d. unde:f we>atkly willne: concljliO:liIs~ tha pirCSCI'lCc of ~emcli!LI:m ~O.I:Ji could Iil~ oo!'}efi<:iO'!lI~o n:~ gwwtb if ~her.e is ".p(lO~ ' supp.!y of [pQtassi.1lI.nfI.,c., the pa:maliN!,placcment i gfiOdID~m bYFota!~U1m.i~im!VQ]yed i~tlUi p:niMern. 11te: ]968 I@Xpelim,e:Nt LifJdllded stlllldil~s 01'1 the cff!!,ct£1af8() d:iffcront. killds of saU, 0.1'1 rice ,growth., <m.d b) pOltii.ls~!LIm SUlPiP:.ly em [.ice growi!h ~n the fl'eSelllJCI'l Ow !lbs~oe o:fa .m.adiafate CO:lilCClatrniliO:ITh of sodiiill!:m. dllioridc:.
Etfects ofl ~ii!!I!i!l!~ ~Ichlifn. i!l!1!d '!In~;g~~ilJlm d1~Ql:'ides ,onr,ice 'lIflOW1h The ~~~oilcult:IlI:fi:l Illil:clruf.!ru~l:Ie 1Ji:'l~dI.iil1l.l~i~'ar"~ c:X!~H~',r,ime.n~, waS" used k:I 'Ij,tudy the effec~o~ oodilJliD. mlillgo~S'lulm.an(l ca!,cilJ!tIoh~ioridcs 0,[1 d~ growth (F~,g..30)10 The I~OO>!.icity hi~ CO~C:'I1)Jn,tro'of l~oflsof lC'aillcilum.ch~owiide appeairs~ob<e: a HtUcles~ thillll thai ,of so(ijlllm Mid magtlcslJ;lm. dhilloddc:s. The pa:nide wej~l:t of th~ plamln Wf'lI~hl'! ~()di~UJm ~d: ~na:gnils[1Iam :ii'Il;l'IDe~ wilisooducelll ~!O ~ilIiIrth,u, of the Oo.~lrQJ :~tOi!bgu:t 7.,lOOui ~QrI~o:s.cml~l,
IPt)W$iilillliQl

WeeKs cHew '"[t\Ofi5p!enlilng f-lr. :;la. TUlctlnt p~r.rQrJj;wi'i¢c Qf ~l!X~citi~s gn,lll'i'n illli ,cuU!li!~~S'DhnioJ!l.

kiw-· ~d hii8h·tm~dtl8

ocJI----I-----i-----i-----i--r-i-r-il

.L,

--I---I--;-~~

Wt~~

O!!i'!!tflt 01 WlGr 1lIOlI~ ,t"Iol

If-liS. :2:9. R.c.Iii:UQ1'l!~njJ,Pb~Me~;m R.1:R ,~I1,d .nBw.G~n. onll1iIM c

Oifle>l1 bl~d¢5QrU'l"o~,~rii.~H~·s:. Ptlt;'l$$ilUmtl

feQlilitememt of~e

!fiee pmilmt will 'd'!·e

abm!~ Of presel!lcuof s:edii!l!m'!,l~I_:ide: Th(!pl;.m:t~ W~~~(l'wn at difr~:ro:ltt, potassium ]evels iill the :prescncc erabseaee of I ,,000 Pl'In N;JiC~,

dcf[~el:l.c.y' s,ympi~oms start!!,~ til' ~1:'UI~lPJru:JtiJ1:g lai ~ow po~'a:Ssi'U,ml~\I'i!]s (1 8TtI.d 2p,PJml K). Lates, UlC p)antswilllh :5 aadI 0 ppm. K .aiIll0 d!eve~~opedls]i8ht ~ymptQm$ of [IlQt:lSSi.lIUifiI. defi.clcflCY.IiiIth~ a'~~,~ ,of s:od,illm c.w.OIridie. o(J'omge.yeUowish dis:co]omUo:I1 Sh!fll:ed wHih tllile .I.ow,erle~v;e$ from the tip$ gQ.[~;g d)mWlw.a~{I, Tbeleilvlls beea:I:I1.cd:mo,!)y" wHh SOlne' OftJite\liIl silQwir:l,g, a Ji~w brown SPQt~ .<li!!I(I Qtheri drywg IJp.[n tiliIe proSfl:rlo(J of scuilrul1tncNoridill',.tIw: leaves remme,d erec~,. wete:lJiI'il'e!"!e~. :tn,d dJjd rn,Q~
a"e:ar ;a,bolilt ] rIlQJ[tnafler

hai\l\~lbrown :iipotsll(f~g.

:3:1).

lS

o No CI2 •
\1

Mg CI2

Co Ci;"

oL_~~~----~----~--~--~----~
a
2000 4000 6000 6000

variety, one would not confuse iron toxicity and potassium deficiency symptoms. As shown in Table 9 and Fig. 32, sodium can partially replace potassium at low potassium levels. This seems important in weakly saline areas where frequently the sodium content of the plant is high. The presence of sodium chloride decreased the potassium content of the plant when there was a high supply of potassium. It appears that antagonism between sodium and potassium in the rice plant occurs only when the potassium supply is relatively high. When the supply is limited, sodium can partially replace potassium. With these considerations, the relatively high sodium content of the plant can be beneficial to rice nutrition under weakly saline conditions.
Zinc deflciency of the rice plant

K>QOO

Electric Conductance of Soil Solution (I" mhos em-I)

Fig. 30. Effects of sodium, calcium, and magnesium chlorides on panicle weight.

Generally, when the potassium content was less than I percent, typical, easily identifiable symptoms of potassium deficiency developed. It is interesting to note that IRS does not develop tiny brown spots on its leaves similar to those caused by iron toxicity even when potassium deficiency becomes severe. Although some reports say that the appearance of small brown spots is characteristic of potassium deficiency, this has not been the case with IRS. Therefore, with this
Table 9. Effects of a moderate concentration rice plant. Dry weight Potassium supply (ppm) shoot (9) No NaCI 9.0 1!).0 24.2 31.7 33.4 39.9 47.7 1,000 ppm Nael 15.7 21.8 30.4 38.5 36.3 36.7 44.2 No Nael 0.33 0.56 0.76 1.23 1.18 1.51 2.75

Observations of the visible symptoms on the rice plant and the results of simple greenhouse experiments have shown that zinc deficiency occurs on calcareous soils at Pant Nagar, Uttar Pradesh, India, and at Kala Shah Kaku, Lahore, West Pakistan (1967 Annual Report). In 1968, the work on zinc was continued by (1) conducting greenhouse experiments on the occurrence of zinc deficiency and how to cure it, (2) studying some factors affecting zinc uptake by the rice plant, and (3) conducting a field experiment at the Kala Shah Kaku Rice Experiment Station to confirm the existence of zinc deficiency and assess several methods of zinc application.

of sodium chloride on the growth and potassium requirements of the Nutrient K 1,000 ppm NaCI 0.30 0.56 0.77 0.92 1.11 1.34 1.80 No NaCI 0.14 0.15 0.12 0.10 0.12 0..09 0.10 content of shoot (%) Na 1,000 ppm Nael 1.63 1.76 1.39 1.24 1.37 1.03 0.98

1 2 5 10 50 100 200

36

Plant Physiology

'(iU

availability of zinc and/or the uptake of zinc by the rice plant but also other factors retarding plant growth. These results were confirmed by several repeated experiments.
Comparison of methods of zinc application

Fig. 31. Effects of sodium chloride on rice growth at low supply of potassium.

f.
~20~--~~~--~-----+-IJ)

Potassium Coolenl of Leaves ('¥o)

Fig. 32. Relationship between potassium contents of leaves and shoot weight in the absence and presence of
NaCi.

Effects of cellulose on zinc deficiency

When zinc deficiency was reproduced several times in the greenhouse using soil from the Kala Shah Kaku station, the soil, without added zinc, did not consistently produce severe symptoms of zinc deficiency. Field experience has shown that the disorder is usually found in low-lying patches, and that drainage alleviates the problem. This suggests that the disorder is associated with highly reducing conditions in submerged soils, and that, the addition of organic matter may aggravate it. Table 10 and Fig. 33 show that (1) the addition of cellulose to the problem soil aggravated the disorder and markedly retarded plant growth and (2) the addition of zinc counteracted partially the effects of the cellulose. These results indicate that the application of cellulose affected not only the

Different methods of zinc application were compared to select those that are easy and safe to use and inexpensive. Cellulose was added to all the pots to make certain that zinc deficiency was present. The control plants (no zinc added) started showing characteristic zinc deficiency symptoms about 2 weeks after transplanting. Three weeks after transplanting, the plants which received zinc either in the nursery bed, in the pots after transplanting or those dipped in ZnO suspension before transplanting started showing slight zinc deficiency symptoms in their lower leaves. The presence of cellulose caused the zinc deficiency symptom to persist. The dry weight and zinc content data shown in Table 11 indicate that all the methods tested were highly effective in correcting the deficiency. The advantage of the nursery bed application is that it eliminates the possibility of zinc salt affecting microbiological or chemical changes in the soil through which plant growth might be improved. A field experiment was conducted at the Kala Shah Kaku Rice Farm in Lahore, West Pakistan, to verify the results of the greenhouse experiments and to evaluate different methods of zinc application. The experiment consisted of 16 treatments, including nursery bed application, mainfield

Fig. 33. Effects of cellulose on appearance of zinc deficiency of the rice plant (O.M. stands for cellulose).

Plant Physiology

37

Table 10. Effects of cellulose on the appearance of zinc deficiency, dry weight, and zinc content on Kala Shah Kaku soil*

Treatment Zn (ppm) 0 10 0 10 Cellulose

(%)

Dry weight (g/pot)

Zn content (ppm)

Symptoms

0
0 0.5 0.5

7.7 12.7 2.2 5.9

14 27 10 20

Moderate None Very severe Very slight

* The plants were harvested 5 weeks after transplanting.

Table II. Effects of methods of line application on the appearance of zinc deficiency, dry weight, and zinc content on Kala Shah Kaku soil."

Method of zinc application

Zinc applied (ppm) Nu rsery 0 100 0 0 Main pot 0 0 10 0

Dry weight (g/pot)

Zn content (ppm)

Symptoms

None Nursery application Main-pot appl icat ion Dipping of seedlings in ZnO suspension

0.8 3.1 2.9 3.4

9 12 22 13

Very severe Very slight Very slight Very slight

The plants were harvested 4 weeks after transplanting.

application, dipping of seedlings in 1 percent ZnO suspension, foliar spray, and topdressing at the time of symptom appearance (Table 12). Zinc deficiency symptoms were observed in the nursery bed when no zinc sulfate was added. The effects of zinc application on rice growth became apparent about 2 weeks after transplanting. Foliar application and topdressing of zinc sulfate caused a remarkable recovery in plant growth within a week following application. Grain yield data showed a striking difference between the no-zinc plots and those in which zinc was applied. All the methods of zinc application tested proved effective, resulting in grain yield increases of 1 to 2.5 ton/ha. When the occurrence of zinc

deficiency can be anticipated, the dipping of seedlings in ZnO suspension is the most economical. The amount of ZnO required by this treatmen t is I 00 g/ha, which costs only 12 US cents. The nursery bed application is also highly economical. Assuming that the area required for the nursery bed is about one-twentieth that of the main field, the rate of 100 kg/ha Zn in the nursery bed corresponds to 5 kg/ha Zn for the main field.

Factors affecting zinc uptake by the rice plant

Total and available zinc content of the soil. Neither the total zinc nor the available zinc content of the soil was related to the plant zinc

38

Plant Physiology

content '(TflIb~e13). :S!L!r~.iS~fI,GI,y.lh~PfQbIJJlll oils cro'a:scd wIDth ~l1ic[~~ing (;O!1ce~tii'!lt:iOilil:S ofl'l,il~getl s a;ppeaJed to c:()!nlil,in <til! 1Jidequa:w amount of z:i!l~. lll!tb¢ ,~ti.ltlU~ soiulioil1i (Table :14). O~villol1~ly the W',eater supply ot .rnilwogen PIUP1IDtod p~atI1.~ growth. TII.i:s suggests that w!llt'l~ plant growfh :is, i;mPJ(lv~d Effer.;IS ()ll,iiu(fgt,~. irorormate ion eonce:mra· b tf'm~. ond pfl of ,~~ltuJ';fl',:'>oiU1!iOJlS on ZInc-oJ as a re~uh offe~tiruizaHOII, the ~i[Jt; [~quiretm[l~ IQf me lice. plaat i~,!,l)$O wllicreasc(l" 'I~us ,req'J;d~¢!J!eJ[t tJPt(J,ketmd d:istrilml,{vu iI~ therke pianl. $ir!ce the mOlY ~ot b~l'Il!ilt Illy so!ilis wil:h ~~W lwailil;abllc zi:nJc: di:r.o:r<l:i,er has ~epolf~e:d!y ~c>Cli)mem:()!rc"ppalle!I'I't with (h~i$ltrod,IJIICHQ:n of im~,~:rovedam!ties which IOOl'!tent TIM;:!]JH@Il~Y s1ilighUyaifec:ted th.e,aib:'lo~p¥ arc: he<1lvily f~rtUil!t:;d. the df~!C~liof :uaj~ro£le;1l Her! of ZJ:nIc-65,as l.nei!~ilireld by cOIno()'ll"lrat]OIl or IllJtritian 01'1 zinc, uptilk,c by tI'~e rice [l!~a!!I~ W~~¢ by~ot!lil !.!pt*c. H~gh ,coni)~nna;tkml'i. of tha bic;ar· sil,dj,.e:di Ulsi:nll ~diloacti¥e ziac, The ~:rf~(;ts of ~ollau [QfI, hOWJ!!;vC!r ,I:'C4:11aril:aM,y aU1il~ed fhe ~ ~rilbu;l]On of lmc-6:5w ilIe reets and. top~. A much ~ic"Jrbo~ah~ ~on. ~!ld [:I,[-i w~~ s:i:lJldi<l!cli beeause abo, hjghcol1~!iltmtiO:lu of b]ca~bonate kmin SiI.!b- higher re~01!l]~a!1ll0u:llJt of Zin:c-65 W~$ rO!l!!!d .i:n. the m~~,g,,,,d :soUsreslJrut from adC[ilio:~s Qf ,o!'gIlllic plIUrit fOOts ~olhe pre:se:nloe Qf SO m.M. bica:r~Olilate ion than in ~e case of!lil1Y oth!erlre,a tmell:t. Thils rna.ttlf;~. F~'fll,ermQ]ie" the ,pJo'bmem soils \W'le
,aJk:ailli:n~.
ilbiSOl[t)UOI1 of Zrn,C"65 bgthmcQl1oent~~il:iWi,~di:l!l iJTImobiJ~t](lri of zi'll!c 1[1 ~·roQit!S m!!)1 be one

ot

The

bylhe

!:iccp,]afll. ~ota] lJ:ptak,e, il]'

th~reilsQ[ls, why lhe !!!!ppH~atie:~ of cel]~loi¢ a,ggrava.tes. zinc defi~~e.l)!cy.

Org,mic
PIQ~
IFIO.

(3!r~in
y~i,lld (ltrglha~

Fielldl Solrface· ,appUed

m.!!iUe-f

(t.Qn/h;i:ll'

~
2

0
0

.0
5
{I

4315598,2

0
'10 110 100

4'm9
fil47 651115
68:55 52:53

3
4 5

'6 i S 9' 10 11' 12··"·'

13

0 0 0 0
'100

10

100 HlO 10 '10 100 Dlipp,tng seedUng~ if I, 1%ZrlO sY$pen:sifon

IFoli;TI'~pr.~v (1 0 kgfh<li~'

'100 0 0

o o

5 5

4S6i!l
6170

.5 '0 .0
110
100

6020
'!'l658

14

15'·'·'
Uf""·'

o o o e

o o o

5713 0008 6915

10 100

15'6190'
!!H70

.' Alillplo,ts were relftiHzied wiilh1!68 '.' •. Appilliedalbol!,!t '2~1l;~

'kgilh.O! N.

67 kg/h!'! p·z0.5,. ,6; I\;g/ha K~ O.

~h~rlnml~phil!'!I~ingi When lil'lie cb:!ihoiefllcy ~mp~om'swere' notJi~.

3$'

Table 13. pH, total and availablezinc content of soils, and plant zinc content. Available Zn" (ppm) 3.4 3.1 4.3 4.3 3.1

Country

Expt. no.

Soil

pH

Total Zn (ppm) 23 75 75 82 110 99 75 92

Plant Zn (ppm) 46 12 8 29 20 10 11 12

India

5 10 13 .. " 15 18

Lateritic Usar Calcareous Regur Lateritic Calcareous Calcareous Calcareous

5.3 10.4 8.3 7.8 5.1 7.9 8.6 8.6

Pakistan

2t 6 7

.. 0.1 HC 1 extractable.

*"" Pant Nagar, India.

t Kala Shah Kaku, West Pakistan.

Table 14. The effects of pH, nitrogen, and bicarbonate ion concentrations of culture solutions 011 the absorption of Zn·65 by the rice plant." Nitrogen (ppm) pH 5 20 40 80 pH 8 20 40 80 40 40 0 0 0 10 50 18.3 38.0 54.3 38.2 39.4 4.0 6.0 6.6 8.3 6.3 705 1069 1350 1199 698 17790 20970 16510 24360 12.9 40.6 73.3 45.8 27.5 (28) (35) (25) (15) 106.8 134.4 137,0 153.5 (72) (65) (75) (85) 147.4 211.7 182.8 181.0 0 0 0 19.3 37.5 48.7 4.2 6.2 5.3 581 1376 1778 8037 13250 26950 11.2 (25)t 51.6 (39) 86.6 (38) 33.8 (75) 82.2 (61) 142.9 (62) 45.0 133.8 229.5 Bicarbonate * .. Dry weight (g!pot) (m.M) Top Roots
cpm/O.l

g dry matter Roots

Total cpmx 103!plant Top Roots

Total

Top

.. Measurement of radioactivity

was made 6 hours after Zn·65 was added to culture solutions.

** Bicarbonate was added as NaHC03.

Figures in parentheses show the relative amount of radioactivity

on the top and root portions.

40

Plant Physiology

,Mobil'i.ry of Zrn,,,,,(j<5 aOs-crbed :v.!Qf1Q().r'S· W'~d,~. tower lea( act the .rice plrm.,~ Ta,b]c .1:5 showstlfuc m;obiility and ,W!itribllit~OJJl. ,of Zin:o.{iS wl1en iit j~ ;JJbsorbed Ul~llughl:heWQtso~ through.the:5:til. ]eaf from th~ '~QP.Wh(lml .zm.c w.as ilibl5:O!l'ibedwol:lgh dle:
I)oots,. a disHm:::U\\, highe. amou:n.t

'hbl~

.I:~. MOib'j'ni~li'IIJIId ~.iMlI'i!mIlilon ,1h:roQgfrl ~h~r@Ob,~iidi ~hwl!Jsl~ :~ I(tUiF.

Qr

lrH:iS~bwJl~d

went

to the
~I 47415

u[}per[eaves. A sini!,!l)fh'\e[Jijl\wa~ (ilso :foulf.!d\vh~~ z:mc WlJ~ !l~W<lil1mdthrougJ~lwl! :~o"iWsl~.eaf. These: eXpeil'i:m.el:l~lS ~nOliyex~llai:n wllIy bl:'irwn d~~!Olg.!A!~i,o:n $W;rll wjtl~ tllil.e ~iO\W], ],eaves,

2144

3 4
5

985
6159
!'il1:;!!5

'5'1 92 2039 118~4

509 sns
lUll 9'19 .:198

Pan~cle emerge!llce

BrI·16 •. !!.pho[,Qp!!r1od,.sensitiive va:rrety, ['.eql:iU'c'S <! mil1JillnUm or ,eightpho~DL~d!i:!cHve tyd~s Ibefore ~ili!ljdein:iti:!tio!1i calll! ,i;lCCl.lf.A!~(]UiQl'la!I :s,ho:rt ~hotopcri.ods OIi~mcccss!liry for the p.anicIDe pri~ mo:udi,(lm to develop fi!l!rl:he~and ·emerge :O~~fwise, the p~~or(Ul.lm will :l'eVi!U toi~S\i'cget3ti.ve
eharaeie •.,becQI'I1I,e dQml<lfltWlid ~:F(ld,UiCe !J\I'ege1:1l!live braneh, olf wil:l d~'V¢IOipl@l.O a sm.alm pan]cl.e but will Jll.wer ex:sutf~am the nag 1t!!!f~h~~th, Otu:erv1llti0J18 hOli¥eiJ1!~cat~d f:I defimw ,effect ,of

~O!DO

3
.4

435 S4~· 269' 1169311

5139 U1200

:Pw~(l'tQperiQ~ em [hi:! slll:lsequeilll de,vclo'pmenl !of thc p,mi,clc.. 1'0 stlLuiliy (l!js ,eJrec[., 1lfl~xpcrlmernt
""'liS

conducted usLIl:@;DIP[~ 76.

'ID'jjjb~ W6. Hf~cc~ o:rp~~g~il;J;lC'l'iud~~:b:nlp~,j:d~

(!t~r~

o:f ~(lpLm(s),

Gmc~iU!:'ii

T]u,e;p~<!:n:~~we~'e ~O.wlil fO;I" 30dl<liY,S! undc!r a 2i4~.Q~lf ph;QloiPeri:o~ (nof!,wld:pc!i.VC: IPJrlot(lp~!Ciod) fi'IJ'u!rfiber at U)hr WIld ffie:l'l gJ\J\e.1l IO;mel 15 li!!·~~hOl.l~.p]lot,O.illldUictive cycles. A:fk:rth~ phO:loifld!1,!ct:ive ilr~atm~I1U.the cycles t!!'i'O :~,ts of P.llmits were (ltvidad WIl~O :1>1:\ gmlil[l$ .p.c:r ®'t <!!IiI.:! ~I'I [0, 12. ~I •. [4., U.sand 24 hOl!U 3 P'li1!Qtiopet.iod.Ll:nti~n'O\lrerilin~. i 10 As sbOWIl on TaMe 16, aill[pJants given p!hot;o. mdluc;tj\1\ecyCi.es iIlIDt:ia~:CJdi }3fI:icle p'I'ilimQu):iummdi 1

Pllato,period "f~r indw;:. ~lve cyc~QS ~.hourd 10 12

Daty:~ '~mm Leaf lrej3I~m.e!flt number of

tQf1to~,r·
li~!J

flowerill!:!
12

or

13
14 1Gi

39 53 60
6:2

~2 ~2
ii

,eXSl'lrUot!.s,e)tce,r ilt '{hl':: :24!-hQ'tlt t]a,roc' lil]aflJlS OUi~ of .~ did .nOlI 0 exsett paik;lc:s:. The IOtlgll:rlhe pha:tope.]Qd! 1Ifhl',r .pru1.id0 matja,tlOln. tllit~ .g~21:c,r \VilS the delay in !I1'.io~e exserlilO[I.
~iltto~lOplu.i,~iwhc.1iII

od!I had puJde

2~
10

67
70'

12 12 12 12

t2:
U3;
M 16i

4~
46

38

~2
12 12 '12 '12

'When. thephmts

tha!!lthe mimWllI!l;flll pim:omperi.oo:s ha.d.\I'crylwU~e

~m giVi!iIlI flvc cyc~e,s more .(CQ!l:Ii:f(lmc!'t:t. Ute SIii:bs~qllc~1

~7
46 46

·efrec~

011 pmi,c;lc

ex~erti.oJ~.althou~ th.OiiiCsllbffie~,~Jl!IIto the 1000holJ!ii" aad lZ·lI!o:ur l'l;no~operilo~f](J,we:f(lldl si.iJiil):iU!t~y ~ A:'OI1eri!l~ G··f' ::ieveli'llplants. cad:icw' In:alD U1.eollil.crlRa:tl1lle:n:ls;. Appall'elltLy. tlre L$!rrl! (J)'~I.ifor ~0 !cycles '" 3.8 ~ve e;x:t:ra cy';]~$i)jre ()~!crulgh~o p:f(ldl,llc~ the ~Ie'ce~ sUjmu~(ls fOJiIDe mo.nnarn e:lo:ng:atioll 0:[ LSiEl, (.0I1If1o, 11 i5, c~cle'5,.. 1;.5
1

:2~

4.~'

Table 17_ Effect of photoperiods after panicle initiation on the lengths of the panicle and the first internode, and on the panicle-internode ratio. These data from 10 plants are given in centimeters,

Photoperiods Treatment Panicle 10 Photoinductive cycles

lnternode

(hours) 16 47 20 2.35 29 40 0.73 24 51 .. 15 " 3.40 26 38 0.68

10 23 30 0.76 23 30 0.77

12 31 35 0.89 28 37 0.76

13 43 26 1.65 29 37 0.78

14 44 27 1.63 29 38 0.70

P/I ratio Panicle Internode P/I ratio

15 Photoinductive cycles

• Average of seven plants. 10 Photoinductive cycles LSD (.05) for panicle == 5.3 LSD (.05) for Ist internode= 15 Photoinductive cycles LSD (.05) for panicle == 1.2 LSD (.05) for 1st internode"

6.6

2.4

the internodes and the eventual exsertion of the panicle. The panicle primordia of the treated plants were all initiated at the same time or at the same leaf stage as evidenced by the fact that the leaf number of the main culm at flowering time was the same for all plants. The delay in flowering was mainly a delay in development and elongation. The lengths of the panicle and the first internode were not greatly affected by photoperiod after panicle initiation if the plants were given 15 photoinductive cycles, except at the to-hour photoperiod (Table 17). The plants that received only 10 photoinductive cycles generally showed an increase in panicle length with the increase in photoperiod. The internodes at the 16- and 24hour photo periods were shorter than those at the 10- to 14-hour photoperiods. This shows the effect of photoperiod after panicle initiation. Whether the effect of photoperiod is directly related to internode elongation and particle length or indirectly to the degree of development of

panicle primordium, which then controls the elongation of the panicle and internodes, is not clear. The degree of development of the panicle primordium may be the factor controlling the length of the panicle and that of the first internode. The panicle length-internode length ratio tended to be higher with longer photoperiods in the case of plants receiving 10 photoinductive cycles. The present data show that when the panicle was greatly elongated, the first internode suffered a great reduction in length. Some plants grown at a 24-hour photoperiod after the 10 photoinductive cycles did not exsert panicles since the first internode was much shorter than normal or even shorter than the leaf sheath. The increase in panicle length was due to the increase in length of the panicle axis, resulting in a decrease in the grain density of the panicle. Effect of temperature during photoinduct.ion The rice plant may respond simultaneously

to

42

Plant Physiology

te;mpc'ra;LU~eanld phQito'~:riod but th.e de~,~e VlIrL:e:!l

acoQ!Jdingto,va:rielty. 'fel1l!pe[atli!r~ affe:cts both till!! plloilopoeriod~li!!sil~ve·arldphotopel:,icHID·i:Il:~e:~sili,1!'e varieties. Gem:rainy, hliFJh ite:mpernlllfe8J~cel~:r1l!'~es h('!ilding {iindlowtempeil'l!h~re' de~,ay~it
J\l,fi1LmJgh there ~a;\!',~bee'lfa. manYPlIb]'ished
nOW1;lI:~

~U,l:di~~of! the Jeffec!u'of~ern.pe['du:re on tillJC

lilg

OOSpo,i),se: the riceplant, ~e ql.!ie:stiol'l sHU (c'!ll<!if!s 35 to whethe:u t:CIOOlperntu~e's:ffccl:sp11:QI!0"

or

i:l!I:cllllct!Q1!Dor pmlllele in]UatiOill.sill(;(l the: ~!leviOiIll:S studics< l1~v(l bee'n based 0111eitlil~r the growth of ~hjepa IIIide or oln Ihe time of no'w~[iiilg. To ill'l$;wer the ~bcl\~e'qyes~i.o!l1l., anexpeiuirneni was c:(md:ucted IJsl,".g, th~ v;I!'iety BPI, 76. l"~ plants wc:~e g:rown l!llde:.u OJ 2~hour p.ho~operiod for 45 d'il.!ys;:tft~~ which il.~ey wt::~egi~e!l!]4 pi~Qto. indUlcUvc cyt:lci5: fI.OI hO!llrl>'~:lieh) with the follow· iri;g f!igh~ ~em,er<lU;lJCs: 21. 27, and 32 C. 'l"ne
~1;8ifl~SW(l~

IF-----+~j~~~-

.,~I C

Z?'(;

V ~C

:again sUibj:ec[edIO,

th(l 24.hQIlIi~ p.h,o~o.

10 II----I----+---!----I----I

peil'liodliIDfl:cw U:e;~;tmenL. the T!llMe 18 :d:!Q"Wll tllilllt p~al'll:S gi!Ve:.I1. p._hQLof.lILdtlct~ve ,cyde~ ~ttl 'e O(l!w·e"ed 38 dl)y~ I)n~r

3O!
~ ~U~llii:fM f,rom, Sly!!'1 or Tr~OI !iPeti1

t[~!lJtmemll w.1:lli]efilJose ,i¥~Jl photoinduc:li'vc c),c]le;s at :2.~ (: OQwered S:2 d~ys a f~¢r tre!ltmill]i) [or .~,

lfig:.3iS. liffel;'~ @f h:!mr~r~!1U!.!"~5d!Hinl! ,phO'I!!Jind,uclimiion

I~iilfing :ilnt!1if'lj',iJjls,

., 21(;
ZOO 0 211C'---+----+-,f-----!i 'tl 32C

,dliffrl\~"~~ of 14 d!llY~;At 32 therewas also II ,~jelay :ill t]:oweJ'ilIl,g when OOl:t!Jpar-ea '~O d1,e. 21 C tU~1"lt'~l'e'.n t whicl1 WIl5 the o']Hi fill.! [11. Sl:llice [be 1:i!<aflnllu:nberQ!l: the I1lainCIJlm of pJ1.!nfsill Oil] ~realm!e'.IiI~s WiI:5ltibe S9iltne be fo~,e:(l:n,ti,:U l1!owerulI,S. lhcpl:a.m.lt:'l, must ha:ve heeLI'l photo'tC.iIl-

~ndlLlced at tile ~ruH'e'time .roga:rdlcss, oflhCl

~120 ~----_4~~~~~~~~~~I~~~~
£

pe:ra,u:m~ d:L1 ri,l1lg ,pE1!Otoi!flduc:~ ion. Tbe ve,~,e~l<!tWJ!l

c.Q!lVe r~ed to rep~Qdtlcl:iveprj. ~'m)1d!ia,a l d~c same thne orfjJt the same morphologi~;llIU<!g,eJeg<!rdle~ o:f~ernpcralLlr~.

"" ~
idfIl

pril:nQi.rd~!! We~,e,

;Q

m
];;

if aD

~---___4---~~~~~II~~~""iI

"fhe su,bseqJllwnl
Il1tC~t$.

!di~\!,(!IQlp:I!!l~!n.(lIliIid eloogatiolWO(

'!h:e p!ll'li!~le pl:im()rdi~ ~iff~[¢d in tile lh~ee rreatwitlh h~ghtempera:t:u'ue8i!::~.lc":liting the

prooe'ss (Rg. 34 ).leaf d(lv;el'opmel:ll W<lS li!1~Q a'CJJD~~ctlilt.edby lii:jghlCm:peraU.!~es ffig.35).

'1)IlHj.mumpa,1iI ic]e dCll'e~,opmcrn:tandleaf
l'!l1e:~1

CI~velop.

Q.f tbe mi!l~t'l cl,ii~m eecarred at 7.7 C. The

00y5rrom :Flig. ,34. :Effce! Oflc.mp~rallllllfes: P<J!iilid¢ d~\I'it;llo:p,mel1ll"

Smt'l ,of TfeOll1l\43nl

d!u,riiigphQI!!Jindll~tiMl~"

d~.layi!"l p'<!lliclc e~~euio,1:I of p]SifllS ,given photoilld'I,ICliw cyde'sat 2] C ,~:Jil!d2 C Wall m<!il]~,y 3 c"lJ~d by Uue $~:Qwlc.afiJl!lilllef'l'at A1lhol:!g:h the !;l~:a:Jl~; \1!!Cfe 'UifH:ler thesam.e tel'Uiperafi,l re (;O'Jld:ilkni$dg.r.iflg dll~period of

43

Table 18. Effect of temperature during photoinduction. Days from Temperature treatment flowering 52 38 44 to Leaf number at flowering (main culm) 14.0

21 C 27 C 32 C

14.0
14.0

panicle development and elongation, those that had been subjected to high temperatures during photoinduction (14 short days) had a more rapid panicle growth. The effect of the temperature treatment was apparent in the growth of the panicle and leaves long after completion of the treatment. This study would indicate that temperature during the dark period has little effect on photoind u ction or panicle ini tia tion. However, it a ffects the subsequent development of the panicle p rimordi urn.

around Laguna de Bay, the water depth may reach I meter for several weeks. Varieties used in these areas should have a good plant type and still retain the ability to elongate rapidly if necessary. Several crosses have been made by the Varietal Improvement Department using floating rice varieties from Thailand and the short, stiff-strawed varieties. The, response of the parents and selected lines to increasing water depth were studied. The seedlings were sown in clay pots containing soil and su fflcien t fertilizer. Thirty -five days after tran splan ting, the plants were transferred to a water tank and the water level was adjusted to J 0 ern above the soil level. The water level was increased by iO cm every fifth day until the water depth was 110 cm. The increase in culm length as a result of the treatment is shown in Table 19. Among the parents, CP231 x SLO·17 showed the least reo sponse while both floating varieties gave a large response. Selections from crosses, with CP231 x SLO·17 as one of the parents, gave very low responses, i.e., these lines may not do well under deep water conditions. Crosses of Leb Mue Nahng with Taichung (N) I or JR95 showed the greatest response. In some cases the response was greater than that of the floating parent variety. Although the water depth used was just over a meter, the test showed that it is possible to breed for short, stiff-strawed varieties which still retain the ability to elongate under deep water conditions. Apparently, the height of the plant and the ability to elongate are separate characters of the plant,

Internode

Elongation

Response to increasing water depth Large areas in Thailand and Pakistan are planted to floating rice. The water in these areas is usually more than I meter deep and remains at a high level for long periods In the Philippines, as in areas

Table 19. Response of selected varieties and lines to increasing water. The relative heights were obtained from the Varietal Improvement Department. Variety line Parents CP231 x SLO·17 Pingaew 56 Selected Lines I R435·3 IR435·14 I R435·17 I R435·28 Interm. Interm. Interm. Interm. 97 91 111 92 119 108 127 113
Continued

and

Relative height cI assif icati on

Control

Treated

Difference

Short Tall

91 164

93 196

2 32

22 17 16

21
on the next
page

44

Plant Physiology

VarietV;;'lnd II'i~le IP!Jnm~

IRe:l:ati\!!l ~eight
cl.ii!~Hf;(;ati()!'1I

Il..ebMue Na'hi'ilg
1<I'UCnUlr1,g fN) 11

TilIlll Shor~:

1II';:l:7 73 76

2'15 107 "105

48: 34
30 23: 26, 20

Se l:eClledlLin;es
I R439· 11·37· 1 I R43g. ~:44"1

In~e:rm"
lntarm,

5S'
!!lS' 86,

<82
B~

IR439·2· 2-1 1R43!l 2· 7-1 Patent'S 'CP231x: sto-rz

l'-ell lMueNahrlg Selected I.rnE!5 I R44ll-2·2
1,R:,44~·5

SI1ion"
lntarm,

1106

Shon
TaU
Il1!tllHlI1.

91

1617
:97 1 13
S3 ,

93
2115 1 ~14 137
1(13

2' 48 H5
:2111 :2'1

faU
lalll

IR441·110 IR44H6,
Parij";l!~
'Pe~{2

Shon
Short
I

94

1:09

15

!i:~1IF(N)1

l!Web Mue [f\4~li'In!lll';d

,Si Hl7 33

215,

~OO

39 48
319

;Se:I~:ted Une~;
I A4.:),2·1-11· '1 Shart
Shor't Slilort 1~'Iit.e:ml .• Irlttlerm.

~R442·1·~2·1
IIR4·~t2'·1·2o.,1 ~, .442·1-:;m,-2 R IIIR ~LI:2.~1·3S!-3 ~A 4142"1·6,2;·3
IIR442·2·ltH

rs
58 '01

11:27 ~132 ~18,

:5:8

t29
120

68
87 75 75 81

Int~rm.

~A442·1·'12·2

In1!erm. hUerm.
In'~erm. Intel1in.

'1:34 1 14
'108 13:1

60 63 ~2 48 38
33 50

~ R442·2·32·2:
~IA4!410·:8 P~t~
IR8(6S)

T~III
S'hI,ort lnterm, 5hor~

11:28 64
87

eS

121

157
101:

23
29'
37

la'5·47·2
II~ 11· 2:88;.3-1-2

1 25

38
:3:7

62'

99

45

If

....
e

'11le in t,e:graUd $tudy of the relaUo','~shiJ',b',e Zween the

p:roperlie:s of the rice
and
I'Hltririft(!lJalulJ8
gftlr~l

"mel'its processing,

,(t(JtiI'!g.

this year emphasiud: (a) lhtt ilif]ue1'lce o()! pariJ()mr~g Ol'j' lhe phY3iroch.r!mie:a/ p.ropel'tie:$ of t/~(! rice grain Ql~eve~llltlJ'i,etie'S ,and selectlOI'I'S of fica', ,(b) the .na.ttlr,e ,office hemice.lluJoses aud' ,their ,relation to the alkali test" [e) ,rJht impl<.ovel1:!e'!',tof the prot,eb~ eontent o/,rice ,t'hl'ough CNJSseS be/we'en JR8 cmd six ldg/ll..fJ,olein varieties, and (d') the pity icocilemicllJnalt,ul(! o{.ric,(! protein:;;' A study ~f,lhe melaboltsm of ",,'ot,ein ,by '11,(lr,~et'ie/S differing in
grl1iJl,TJ1'()'teil'!

ievel ~vas b,ridaled.

Effect of Parboiling and Variety on Grain Properties

Parboiling and grain properties

Rough rice of seven varieties and selections differing in amylose content was parboiled to study the effect on the properties of the grain. To parboil, the rough rice was soaked in water until saturation, after which it was steamed and dried slowly. The process had no effect on the IOO-grain weight, protein content, and amylose content (as measured by iodine colorimetry and potentiometric titration) of the brown rice. That the starch granules underwent gelatinization on parboiling was verified by the loss of birefringence under polarized light and by the amorphous x-ray diffraction pattern (Fig. 1) of the flour. Starch corrosion by 2.2N hydrochloric acid was greater in parboiled than in raw rice (Table 1). The intrinsic viscosity of the whole rice powder of the waxy selection, IR253- J 6-1-1-2, did not change following parboiling, thus indicating very little degradation of starch during the process. Parboiling increased the translucency of the grain in all varieties and even the waxy selection, I R253, became translucent. Three tests showed tha t endosperm hardness was increased also. The values for the breaking and crushing hardness, as measured with a Kiya-type tester, were higher for all parboiled samples than for the raw samples. The head rice yields at 4 percent milling (the point at which 4 percent bran by weight has been removed) were at least 99 percent of the total milled rice in all parboiled samples, while the average for the raw rice samples was 75.3 percent. The percentage passing through an 80-mesh screen after 40 seconds in a Wig-L-Bug amalgamator was significantly reduced, thus indicating that parboiled rice has greater resistance to disintegration than raw rice. Parboiling drastically reduced the extractability of protein in all samples by an average of 45 percent. Although this was true for all four protein fractions, the globulin fraction showed the largest reduction - 65 percent. Similar amino acid compositions were found in the residual protein of both raw and parboiled brown rice following serial ex traction of the protein fractions.

A histochemical investigation showed a fusion of adjoining starch granules in parboiled, but not in raw, endosperm. In addition, the protein bodies in parboiled endosperrns were less distinct. Parboiling then, evidently causes the expansion and subsequent cohesion/adhesion of the gelatinized starch and protein particles into a solid matrix. This information would help to explain the improved translucency and increased hardness of the grain and the reduced extractability of protein following parboiling. TIle cooking and pasting properties of the seven raw and parboiled samples were also compared. The cooking water of the raw samples showed more dissolved solids and a deeper blue color following the addition of iodine than that of the

Table l. A comparison of properties of raw and parboiled rice (mean of seven varier ies).

Property Raw loss of milled-rice flour) Brown rice hardness (K iya Tester) Breaking (kg) Crushing (kg) Head rice yield (% of milled rice) Percentage passing through an 80-mesh sieve after disintegration Cooking water (cook ing test) Solids (%) Blue value (600 mu), absorbance Elongation ratio (cooked/raw) Protein fractions (% of dry wt) Albumin Globulin Prolamin Glutelin Extraction efficiency (%)
(%

Treatment Parboiled

Four-day lintnerization

60.7

77.5

5.4 8.9 75.3

9.6 15.4 99.8

89.0 9.9 0.31 1.44

48.8 5.5 0.19 1.14

1.00 1.18 0.29 4.95 74.1

0.58 0.41 0.16

2.83

41.8

48

Cereal Chemistry

to

!l!

201

2,5,

IlDif~ro!C1~!i'I Angle

2 ,13'"Idegrees

s.amp~:e!s. 'f,ne 'p3rbOli~le,d grains ~:c!ndl!~dI '~o dxp!lndginhwisc mOl:,e dum d:id tin,era.w .W.ili:ilils. 1I11mS lCl1d:em::y waiS mO[l'1;l "]lp~r!.'llllti!1! Ute sa:rll:p~e~ P~ilQ,~Ik!c::(1nthe cooikillg 1W3e r Ifor 2 hours, beroN! l t h~aling (Fig.
Pl!~bojiled

p1Jifboil~rl IRS.. ni~

~X.p~II!li'liI nile faillllc~

[UrO&l'iIJ(l3~ioR Ul<IY

Mso

iocHn:"e ~;t<liiil'!.i!lg of stereh ~Ili sJiicc$ of p~rbonLl:d JR:8; riceas loonu.parcd wi~h t:i~~.t
of U~eraw ~~~'"

:n.

Pl'Iiboi limlj.l inn uenced

U~e il.1'IlI:yIOigr.rpi,h, C!l!rves of

or

,~UI I ~"iPeroerlililqucoliIs SI!lS"p2IIll:>ioJtlof brown dee pewder, l'lle gclal,iJ!l i~<lliofl lemptmlu,I,res of lhJ:l r.i!lllrIp]es waHl 2,0 pCfOO:IlJ~.,(W3XY}8Jfld 7.5 per{."!C:lu
<IIln)'

The ch~ng,~s i:n :grain propel!'~:ies t1l1~tn.sslm;ag:e the ~l!iw~itld 1?3:rbo~leds~mpJcs wc~,~ also $T'Il.dicd. 1l.e il!rlijI'logruph, 'visco..,illtlei 'of parboiied elee d.id !'IO:leha:f1gc., wherc'.astho~ (1m tl1Q[CIOeJ1l Uy
~'m.rvcs!ed, r<lws~.ll!1pies ~l:IicrcaJ5.edw]U:1

",g~. nUl

lese we~cshjlllcd '110 IOW(l[values, bu( the ~amri]~s wi U~ hfghier a.my]os~ COJl'~~ :i]M:lwCd nts I,ittIc Ci13t1&C. Thc '~,~,k\l'isCOiS~ty ,e:h<llli,gedi din.sl le-

,allyin(hcMgh!l~l (27%) ,:iJmyl~ ~<!mple, ~R8. tfro.llIrl I JII 0 1:0120 Bra:lbc!11d!er unll£ .. SSiI:t:lp,les wm~ 2-,Y'e<lf~lg:!1Ig~studll' ofroHglii 311Ui liI,lnccll rice "!li(] rk!C' March or fhi!! 11~8l;i~fijjHy. less ~h<lrl 25 pen::clll jUl1l1yl~ ~howedlic~iS: 'chOlnge, TU1!!sc resul ts Ji!1aJy 00 rd91~!cliliO ~he ob!5!:lrv.~'~1 ,ltU situ ret rO',rlldi!!:l~.g!l of ~.Illly!g~e ~Ili whi~h ~.]n~. Th:e en:d~pe~ml'ih,Qwcd an lIiLCg;C'a!S~llg elIi®~mrC~I.ijod:in~e.,bllJe test sho,",,!!'IiI ~1]afiiJt the '~:I];Iy~OI'!:C of :!!r~~ngcl;lUji!il~ the $.huch lind PJot.e~!1!parl.ide5 of Ihc'gelaliilllilCd ~i~'l'·amylo~ J~Cle Sa]1l1pte~ W<lS less foUowiflg parbolHflg 31ft ~ COQki;illg (F~fl, 3),. In ~x'l~~~t;)llMe lhlll11hail of dli! ~OWi!T I)my~ooo l<libOir;!tQ~y·p1lirboi.led ~3.mp!,!3:S.. ad]oi.nililjg sl<lI:cl~ SOlin pies. Therelro,graiCI:at:iofl of theg,elalinbred gra:lll'uies were flll.sed tOgjlt~ll!r thro'llllfl tin.ecllli(lo· <!ruylQ,~" w:i1 il;:11 prreVel'l!IS ltlil $\~Ui:!1g of U~i: $i.pcIm. A p,,~boiled ri,~ !l<'!!mple f['omB~rm:l!, Lh.OWg,!:<l.nuks dll~i11g i~e,atil'lg in; ~heillll11iylogra:pi1l bowl, 'e'¥er, ~mo\?{ed sl:ardili gelalim ization of o.nly ~he surcan tl~il~((!xIP1:Qjilil dl~ s''lIfpmssCld :lmy]ogJiilJm of t3JCCeells, A ,crnss~cl~Q!1 of d~~ cuok!eiln:ae griil~lli.

srarch I:lecame meee ~nsohil:bl.e ill. wale:, ~nn H.s ~.my]015C loomefl~ d it! not change :sj!lflIDficiltltly d!u~rng MO~<li~ fOI ,eitn.cr rawOfpillrbQlu~:ed, sal11ip!'I;!~. TIles-I.! .rCS'I.!U5 arc llil 2:g,c.e~nem wi 11:13prc".~ous.

Raw

Cooked
Row Parboiled

Raw

Cooked
Raw Parboiled

IR253 IR35 CPXSLOl7 PI215936 Palawan ACC9800

'"
•

",

.~,
It

,1,.1'
It.
."

'"

"~I

.,.

tit

"~I

,I.
.,.

.,.
0"

.,. I II."

fe'

"(
•••

IRS

". ,.,
a

"""

III III

r rr
.11 ,.,

• " , ,c ",
'" ) C)

'((

"1

'"
fa

t '"

,.

Fig. 2. Effect of

2-hour presoaking on the swelling properties of

wand parboiled rices.

showed that the endosperm contents were very disorganized and cell walls were difficult to discern.
Variety and grain properties

Waxy rices have a compact arrangement of compound starch granules in the endosperm, whereas non waxy rices that are somewhat opaque have a loose arrangement of the starch granules. The endosperm of a breeding line containing 8 percent amylose was examined histologically since it showed a tombstone-white appearance intermediate between that of waxy and nonwaxy endosperms. The histological examination revealed a compact arrangement of the starch granules. Presumably, the cause of opacity in these lowamylose and waxy endosperrns is related to the low content of amylose and not to a loose packing of the starch granules. To explain the varietal differences in elongation during the cooking of presoaked milled rice, a

histological examination of the endosperm of selected varie ties was undertaken. No ge ne ral differences could be detected between those varieties which greatly expand lengthwise and those which predominantly expand girth wise during cooking. The properties of the starch fractions and arninogram of an opaque, nonwaxy (30% amylose) Thai variety, Khao Long 258-22-33, were determined. The amylopectin and amylose fractions showed properties similar to those of other nonwaxy samples. The lysine content of this variety was 4.04 percent of the brown rice protein and it had a normal prote in leve I of 6. I 2 percen t. No unusually high-lysine rice sample has yet been iden ti fled. Varietal screenings were made on samples obtained from Bulgaria and South Korea. The milled rice of three varieties from Plovdiv, Bulgaria, had 7.0 to 9.8 percent protein and 24.3 to 27.3 percent amylose, both on a dry weight

50

Cereal Chemistry

all hllJd low gela,[:i;llli'Zil.tion tempera1L1I,e;6w va detics, Irem Sol.l,tl~ Ko,re'lI,ollle wars waxy ,allidl :fOl.!'Ii'were nOlilwa",y, amyl.ose con· tell rs ~apging behw!,iHI 20.5 'to 24.:1 pen:en l. AU fiiw had l,ow~13Li .. iflation lempenli~l:Ire:s. TheiJ prot.ein IcveiSJi!rlged from S.1 to 8.4 pe~~.lu.
nUlls.O:

basis.. Tbey

u:umic acid {pectin. It was de'void ofman!J:Iosc. butshowed a faint r,e:ac:t:iQf! :pdly:phcII:Dliceomjor pounds. The :residll.al bra'lI stm IOOIil'I,ai'lca larg.e ,aJmOllll'lU of arabinose, xylose, ~:JUd a ctD~e. thus g~d indica ting ,iilil ~[Icomplete extraclion of all hemi· ~f:nul,05il'ls. Pall't of da~ ,a,lkali,·eX'Jfacte-d :n:e-rmce[lu,·

A iice IHe'lliIrceUullases
Hemi'ce!III'lo~s, <UT,epol;Y~i!cclulirid.e!, of :hip mo!ecuJarr weight which ·Me· composed ma.iruy QrJrenh)~ untts (karobino.seand. [)..xylose) ail though hexoses (.D-&ii~3!C~.ose. D-glucme. and D-ma.RI1ose) ,iIIre frequclildy rm:l:Ild if.! n~ociauol1 wi,th them. Aside from. ,ccUlllo!IC, the iI.emiceJll.doscs pmbab1y OIUI~ $tiU:l~ the least s~L!tliied ca:r:bOhydr3'1ie f;uu:li:on of Hl~r'l:CC grain. B~'CSil:l~ o:f the:i.r lhilC'\li':J'l distribtiit i.Oon i:n tliIe doe grain, both bra.IlI,"'£)olis'll ,8If.ld.mIDIcd :rice
fractklf'is of
rn

Ri!,

'W'e~

ilis:ed ,as SQi!ir~e'.s ,of li1e:m.irA.

ceUUllos:es fOil" (hiis, stl!l,dy. Hemice].IiI.lloscs, l!'.Xtraeted dl.l.f.ingth.eru_k.<!]~ ~st Qf ~R8 and SPW·7',6rrulled [iCl~S 3re also beilil:g $~udli;ed.
IBran.·poliid11 liIe<micel~ulD:~

,A 0.2 ~:roerl'l [eoowry of the crude wate~·$(lllul;de hClilliceUl:llkl:sc-s was obtarindl fmm. b.ran·iPo~ish by
wii!.re~ extracHO:Ili. lii.eM:il1lg:" and
iii

bl:eiaichiillg.ean:it

treatment,

Fu~tlle.r

p!.!rcifi:c:..atie:n by

O:'oI<rn:rY~~ie

dig-eoS:llOIl of staseh 3.Dl.tI. ll'icbl,(lifO:ilIeetic acid p~,(llci,i = ~;aUorl ofp'.ro'lem: G.a1,t~ O! 6Ql.pe['Oe,n fecovery of a t. puei fi:~d Ii:m~p::u:atio.n co~tajfljing 11% protein, 38%

arabinose" 19% xyl,osti. 27% gabctose,

acld •. and trace a:mmmts of
,gllICOM.

7%li!l'ioflic

f!ecovery 'W3J$ 10.1:2 :pe:rcent of the tio:n of ~his pJcpruratiolll, 'WIDth hyd,[cglUI pcmxidl! did Iii(j,t r;e.S\!Ilt ;jill ,ge~UOf!, AIk,aJ~·$.O:llJllle :1llie:mlliceJhdo:;;es. ere extrac ted w from w;!l,h:r-wa:shed brn!R'·iPoJ:i~~iiI. 'Ihe PJel(,e.Il~liI1ent 'invdhred boill:iing, coom~:IiI!C:' la~ch ~if,'i!sl:i:O!n. with s o:·amyl.aJSie t 55 C, and ~.shjng with e~lilaRol.The a crude extract (26% yf:eld) W'BS 'i!:dfll,Cted M1.h 0.5)"1
sodium hydmxide under niuog;en, neutralized willi'l acetic add, t~e!lJliedi w]ll.h tliWchlor'Oaoeticacid" dj3~YiIe:d. and. lyophilliiZle:c:Il. A 3·~rcent yre]d of fi.brolis. failly !i'lta[c.r·~I'l!lble br-oWill powder was obtailied. The' O\l'eraU extraetlea [rolll lu,an·'Polilh was 0.78 perceat, lhe Dxllia1.ion of tbe h,emice~lu· ~.osr; did not re;slul'f i.Q g..e:liHiQn. The :h.el1l_:iceU~do~e 'p:reP'3:I1lUofiJ~t'I '7% pro,fei[l (as, g'h.llb::lin)'. 33% Fig:.J.

The; o,vera:U bran. The Dldda-

3fabhaose. 30% xy.I.,o,sll.

rnO%g,fI.la!Ciose

.'l1II~

B,%

(iii) ~rbo.i.k.-d., !lnd, (C) ~k~d.

Cro'Sl'

~I:iOR

'of wit:~· em:!Q;speun: VI.) row. S1~:ii1: th~m~,IO:!i:Y mill.

.5.11

lose became insoluble upon neutralization with acetic acid and was therefore discarded. It corresponded to the classical hemicellulose A. The portion which remained in solution corresponded to hemicellulose B. A minor fraction which precipitated upon lowering the pH to 3.7 with trichloroacetic aci d contained mannose. Further characterization work was done on the alkali-soluble bran-polish hemicellulose preparation. The preparation was fractionated in a DEAE cellulose column (borate form, pH 9.2). Gradient elution with increasing borate molarity from 0.01 to 0.65M showed a continuous elution of carbohydrate material between 0.1 and 0.45M. To obtain distinct peaks, the subsequent columns were eluted stepwise with one liter each of O.IM, O.3M, and O.5M borate buffer followed by 0.25N sodium hydroxide. The peaks obtained with 0.3 and 0.5M borate showed tailing and were probably not homogeneous. Only about 70 percent of the carbohydrates applied to the column could be recovered. Uronic acid was evaluated by the carbazole test since only the neutral sugars were recovered from the acid hydrolysates of the fractions in the presence of ion-exchange resins. A rechromatography of the sodium hydroxide peak yielded traces eluted with 0.3 and 0.5M borate in addition to the main sodium hydroxide peak which still contained traces of glucose and uronic acid. The alkali-soluble hemicelluloses and their fractions contained equal amounts of arabinose and xylose, while the water-soluble hemicelluloses contained twice as much arabinose as xylose (Table 2). The 0.25N sodium hydroxide fraction differed from the three borate fractions in that it had a lower arabinose-xylose ratio (0.9), a higher protein content (24%), and showed traces of glucose and uronic acid. This fraction is probably heterogeneous, but was not eluted out of the DEAE column with O.lN sodium hydroxide. Viscosity was measured in Ubbelohde dilution viscometers in 0.05N sodium hydroxide at 25 C at concentrations between 0.1 and 0.4 percent. The sodium hydroxide fraction was more stable in the alkaline solvent (O.IN sodium hydroxide) than were the borate fractions. Only the latter showed a drop in viscosi ty following overn igh t storage. Preliminary studies on the sodium hydroxide

fraction showed that the protein and carbohydrate fractions may be separated by disc electrophoresis in polyacrylamide gel. A study of the action pattern of xylanase and/or arabinosidase on these hemicellulose preparations will provide information on the nature of the arabinans and xylans.

Milled-rice hemicelluloses IR8 milled rice flour was extracted twice with distilled water at 4 C. The combined extracts were heated to 90 C for 35 minutes, treated with bleaching earth, dialyzed, and lyophilized. There was a 66-percent recovery of the crude preparation. Portions of this preparation were subjected to a-amylase hydrolysis followed by a trichloroacetic acid treatment to give an overall recovery of 0.01 % of a purified preparation containing 10.5% protein (Lowry), 9.5% pectin, J 8% arabinose, 10% xylose, 12% galactose, and 40% glucose. It did not stain blue with iodine. Essentially the same method was used for the preparation of the alkali-soluble hemicelluloses of milled rice as that used for bran. The overall recovery of the purified preparation was 0.1 %. The hemicellulose contained 25% arabinose, 23% xylose, 8% galactose, 23% glucose, 7% uronic acid, and 14% protein. The hemicellulose A precipitate contained mostly glucose and traces of arabinose and xylose. The precipitate from the trichloroacetic acid treatment showed the same sugar composition - primarily glucose. All the DEAE fractions of the milled-rice alkali-soluble hemicelluloses had a I: I ratio of arabinose to xylose and contained glucose and protein (Table 2). The carbohydrate recovery averaged 72 percent. Preliminary studies were made on the herniceliulose solubilized during the alkali test on IR8 and BPI-76 milled rice. The resulting supernatant was separated by centrifugation, neutralized with glacial acetic acid, and recentrifuged. The hemicellulose in the neutralized supernatant was precipitated with ethanol. The preparation was subjected to a-amylolysis and trichloroacetic acid treatment, and then dialyzed. A paper chromatography of the hydrolysates revealed mostly arabinose and xylose in a ratio of approximately li l and smaller amounts of glucose and galactose in both samples.

52

Cereal Chemistry

l';:blc 2. Compositio:rl md: i!!I;rillsi:c~isoosi'fY or ,~b:!:aancdl);f D!EAE·~axU!Io~: ,C\iu'o!iiilOOl!:'aph.y.

ff;l~~if)UIi~ Oir'brtl jj'pO'!i~.

i'!nil milledl d~:tIk:uilH;Qh"bl~

h~micell!~IlIO\.."Il~

,eh,u:ed Cla!l'bdh'Y'dJ'a1Je~ Ar,<:!bil!ThO~

:Xyl~

Uroil"lic [~J G01I'I~ClO~ Glw:ose Proteirlla(:~d (ml/g'

Br,im'pgl i~li'l<

0.1M boratle O,3M bor~r~~ t().5M' bora:!:e

34·35,

~. 9
1

la~26 2S:·lS: :9· 1:2 23·30 '51· 13

23·216

3ij

""6

40

4JJ(I

416 3B
3iEi,

20 8 21
9

<1
-+
24
2'4

0
Ul,l!, U;t4
4-

1~12.~'

2.8

$odii~1TI hydl'loxfde

3:2'

1 14,

Mmed rice
rD'

1M

borOllle

113M I:;i>(lrale O.15M ibor,ale (lI.25M SQciium

hydroll(ide

2:8 2:6

27 24
'20,

21
19

B 5 6; 5

13

'~I

22
2(l1

23
3,3:

98
UiQ

11:8

22

13

23

62

Q:f rod~ctioilil of p:mlei!:l O{l!lierrl during mining is I(n"'~r wiw(l:J') [he pil',e"ftl.imf1lg p;l'o~eiJII.level is high (l'!lb!~ <1). l~ would UU.llil$. ;).PPe'.(l:rhalm:llIlch: of the t in 11111 tile cQop~rat]\i'{l: ]iif{¥jcc:~ w~tilil, tli!~ Var1E:llal Illl!c<rclIsed iP',uci~eiini~ Uue miUeCi r]t:fl f~O!ctiO'~,I'iIII~d !:IQt in 'llil~br~$1 aio[l;(!, :1 mp,.m'VC:!lnentOcpar~me!'l~ 0111 :ifl1lpllovem.e!ill of the -Sil~gJC browl'ln.ce ,~fQ.llily~iiW,<1!S, und~rt@ktilnto ~h~pirQlc;inJ ClQnl:enJ, ,of luke. bil'owUQriee .of F ~. F~ • " ilJild f'~pI3r1tS WroJilf!C~05~~ b~tw.e!erl IRS i1i1l1dSUI>: delieflml1nC ~he va~iaibinity of prlOtein oonliefi~ withil:! , II!1Ii!pro,vemleliil'~ tliils' IPlr01tein o~ C~niilelilt O'f: FUoo
MgIl·pwol)ei'lil
protC]f1..

li'i1rie'HiC s wereillfla~yl!)~ii1
J

for

The F~ sc'!!,d'S,fr'QJill f·~pl;l!lbl~l~o~d p'fo,n~~r) cOi!ile~ 15 .~hi~ t as these oflhe liIi~lp(l'o~ef[l paNl1ts (Tllib!e 3). Most IOf!hc:: plli.mt~ ~:vecl
~ re tll1ep.r:oge~y of IFl plaars ~howiml:g:uciati~cl:y h:igh p:m1eilll ,ocm1:ern.ts.JI1i;lit'ld~~ioilil,. ~~aJnb r~Qma

c:ru:(J)e a [p1la:1.icie, U$i~g flijt;a 1Il!l:d, tltI.c :h:i.gh'pwo!)eillil 1[1iI~. I:\e'Sulhs showed th.lil.t tileh)lp (or ,e<l[I~er) grail:!

t~f1ded to haveloW'e~
botsem

prote,iR

.Ieve~$ dHH11 tile

~or .Ia.tc"produccd)g;rllli:n. The ~Tl!aXLmum dilffc~ellcc obt3:il:le~ W.ilh~1iI~ panidl'! was 5.4 per-

tile l(:Jw"'promil!llines showins good ~mhs were~'I'ed. The ~<I!aflce, 361 plaJilt hn:e:s,. were Oi~S!L'!anl~d 1b~ C1U]iS!e of ~~OwlJll'o'teii!fI CiO!il~el~U lor and "lnde:!'h~ra:b·lc]!<lifIU ch"ra:ctcri'~lics. Many of Ilh~' F~ p bi:OWiil rice f~om~he 1,290itidu\l'idu~1 F 3. pb",~~ hcprcsenli ng. 2:S,3f ~ ~lll!'l~ Ii"iCeS dll'S'S~,fied as '';high prot!!.illl'· ~Wil~ ieh h aid, been :!;!i.ved h!;l!d iI$ high II i'rloteiml!~\I\e:I. <IS llheF ~ par.~mu~.p:ti.lJi:lu. M.ost Qflhc F~ U~!L% !lind ~hlU:o.s:t :.Uof U:t~ .~ 3 p~:a~~~ ~ved possessed [he diW<JI'ufliLei~ll d~a:m~~,eriS:li() of ~Et8, Some (]if the HIi!.'I':i!. ;llIow[l~ghi~ pl'o~ei~ c~Q!~te!llt $
(f ~ ;:Hull F4 carY'Opf>~8> ere d\lV3rF ,~iiIIICS w wH:h a

r~w .of

eerrt of ithedry W>eighl. 'The: V<lr.i!:ltal h:11p:wI!{C,uncnl.

IJePSI[[liI'U!!U

C(}Ji!·

dlUct~d yrueld triil~~ oUl 71 ·of ~hiIJ h.igh-l)[Io~.ein s~lec'l]ol1l~liIu~i:liIgthe dwy ,lUld wet s~i1sonsof 1.968. Th~ii!T y:i!eld~ We~~&~!1e~al~y ]owertha!l1 those or IRS, Pela, I RS •. :lJfld I!Iprn·76. l~oweve,r, tnf): protcjn of c<]lI'yopt:es of the:fI~ '!!"ar~e(]e8grow1:lin Ute d~y

s~aro:n. m:ilgc.tlfoom: 7.8 i~iO ~~2.9% wi.rlh. amman level of 10.6%. ~scoll'l!par¢d wi ~l1. 8J)% ;rg~ Ut8. "lX!% forl.R5" !I_rid 6.ml'o fo.rPe:ta. Sirnilar ~e"sli!lhs W"Cre
eb tlliilll!Clrnl:rl'Qrn the \'Wol t.S;-e\liOO(n. t:r"Oip. ~1ilrthe~ b:~~ls wil~h tb.Q:~"~gh-p.rotem 'v,u;iet les which INUl~e]dgb]y

~,ea~ol~.!!bly~(,)<odpl!,'lr.ll~ype. Prote,ill anllllyses IOf If 3 brQVI'IfI llndha_fld-nliilJcd rices COilfi:rmed :prcvrolls rUldil'lgtl 'Ihiill tl1.~ d~groc

phQlto~e!lsillt]~e Welt: disconti:nu:ed srnlC)(\'; hi~lpil'o~n~e s w.ithll.ood plant ly'pe-$ w'Cr.e .mI:!'eadY!.l!\!',ai]abl.e (.oem the ~{l~ller!O be~weel'! ~R8 andt~le$C .hi8h·prole~,fI Ylli[~~~ics.,
~MiOliej]l

53

Table 3. Protein levels of brown rice from crosses between IR8 and six high-protein varieties and of the two parents.

Cross no.

No. of lines

No. of plants

Cross Range

Brown rice protein {% wet basis} IR8 parent High·protein Mean Range Mean Range

parent Mean

I R 1100 (I R8 x Rikuto

Norin 20) 9.8·17.6 6.4·14.6 5.B·15.3 13.3 10.8 10.8

F2 F3 F4 F2 F3 F4

6 46 .. 46 202

a.i-

9.1·11.0 9.9 6.2- 8.6

10.0 8.8 7.8

14.3 8.4·10.0 9.5 12.9

IR1101 (IR8xOmirt39) 1 44 .. 44 208 14.5·16.5 7.G-12.9 6.8·14.3 15.3 10.0 10.0 9.1-11.0 6.3- 8 ..7 7.5· 9.5 10.0 7.5 8.8 8.7· 9.9 10.4·12.9 12.B 9.B 11.7

I R1102 (I R8 x Santo) 6 Fl 32 .. F3

10.014.3 7.0·11.2 7.4·13.0

12.7 8.7 9.4

F4 F2 F3 F4 F2 FJ F4 F2 FJ F4

32

156

9.1·11.0 7.1· 8.6 B.4· 8.5

10.0 8.0 8.4

8.1· 9.9 12.8·13.6

13.8 9.0 13.1

I A 1103 (I R8 x Chow-sunq) 1 47 .. 47 251 14.9·16.9 6.8·12.8 7.2·14.0 Korn) 11.9·17.8 6.9·12.3 5.4·13.8 14.6 10.1 10.3 9.1·11.0 7.4· 9.2 7.8· 8.5 10.0 8.1 8.2 8.7·10.8 11.7·13.2 13.5 9.7 12.2 16.0 9.7 10.1 9.1-11.0 7.0· 8.0 8.0· B.7 10.0 7.7 8.3 7.3· 9.3 9.8·11.6 14.6 8.6 10.8

I R 1104 (I R8 x Crythroceros 8 50 .. 50 260

I R 1105 (I AB x Chok-jve-bi-chal] 8 34 .. 34 213 11.4·18.8 7.5·15.2 7.6·15 ..2 15.1 10.7 11.1 9.1·11.0 7.5· 9.0 8.6·12.3 10.0 8.0 10.1 10.2·12.7 14.2·14.6 15.4 11.7 14.5

" Selected from 614 plants on the basis of high protein content and/or plant type.

The Spies and Chambers method for tryptophan assay gave low values for brown rice, even after defatting. Turbid solutions were obtained, and blank values were high ~ particularly for the red-pericarped samples. Alkaline hydrolysis at 100 C using barium hydroxide for 16 hours and the subsequent work-up resulted in improved values

for tryptophan using the Spies and Chambers method. Bran pigments were destroyed during hydrolysis. A recently developed method for evaluating tryptophan in corn based on the Opienska-Biauth reagent is being tested. This method requires neither a blank for each sample nor a dark reaction. However, it can not be used

54

Cereal Chemistry

iror :red-perica:!lP0d :sample-'s. ,A_mo!il£!l 'lhe oo;ct!nt mn<edtods, d>e'vlIlloped fo:r ly5,in~ ,a~say mcernls ,[1] ,oolorirnetdc assay based on the reaction o;r 2-chl'lno·.3.S.rUniU()p)I:I'idiitnt:l wi'lh the i?p\fiiml'-anIi no Iys,ing r,n:m P lind t ~ subsequea t co1Iori,rnel"ri ,B!S:S8}1 of tile l'e!i~I'Li,llI,g d~[i"'D[hrc. Tl!e <1I,pp~il,;llbi~:ilyolf t~ds l'Iul'lho-d .for nice is b(l~l'lg,
l,c_l,cd.

da&S,ificd as h~glil~ptrncilll prote.il1 rice w~rc exU',act!.ld by pereelauon, Gh,uc· lin fQl1'Ile{l 80% of to ta~ Ittotei,~ i:n brotm ,ilm~. in t~e wlodd collecuen were allil1lyze,d to (Ic'l.e'rl1llmne abOUI 9'1? in mill,ed dee. The lllbu.lm;n~gJobuLin the changes illl protein qmdi~y bioll"gfU abcmt by' an
Three

or t hCIl3ridi£c!i

samp,.te.s had a mellfl protem level of 1,5.10% we,t ba:S:is"whcr,eas ~'h!e rnmed ~ke :h,ad :l4.8% ,md th~ bra n~p'ol.bh had l6.,2%. Tnij-.s;e I,esul tS,iJlOO' in .agree· ment with :p,nwlolls I1ndLl'Igs that ,higb'l~n'ob,~ilfi 'i,ces [ aJOO l~:udcr to mill 3J:1 d have a more 1I1l~f()tlll prot!f'in disttribuHol'I th.m IOW-:PlOtei11 samples. The' (;lit COin tent of UU~bran.pol ish ,nevc-raged23,.0 psrcsnt. Pml:€:in [rae lions ofthe brown and lIIti lied lush-

iocr~arse

in

prot,ei'n leve], TIle

IniJlillg

of hi,gh-

prole'in brownrice

i,n the MeGi,1 i I'Il;iU PWllJfI 8%,

'rnlc'li,OIl was I $% i'n brown and 6% and pm']JtIlj,fl WIIS 5%il11 brawnand

ill nltiUed r~h::e!.

3% In milled!

rec()'II,(,l:1)' bran-poljsh iii oonimsi tio abeut '10% rice. 'Thlis di:SH~bIllHou gJ:Jdii.'lnl[ ill d'le [)~ot'ei'll of for ]ow·!pr'Olt(l~11i samples, It pl'lot,ein <!!iialysi:s. of ~,h~ f~[tctions, ~s renee ted in the ami mJogramof miUed rioe and lru3111'pO:liSl1:!, The two u:mll1oacids most two milHil8; :fracl:iol1s showed that '91% ef tile bra w;n doe: proWi!l1 W1iS ,in lha IniUed :rkoe alild 9"'&:, affect'eel we:re Iys;f.lli~ 1I:liId gh:J~amic acid :BI:',(liI'l~pOl,jsll p~oleiil bad, on th,!!: average. ] 6.0% 8l1l(~ll1iC acld "",:35 iIII th~ b.rllfI·PQ.JiishfracHon. 111C bto,wll rlee

1"l!bl~ 4. Pr'lJ,lgin

prot.ti:lll~arilil~I~&'

~U"!.t'nl~, or

f'J IlfOWQl! mulll hmd·ii'iim~d. ,fires flOm

~',IaIl'lS of

crosses

bet\ll'~,11

mRS 'a:ml six I'Li,gh.

No. oJ
pl~lI'I~

Brow.n 1f11ce

Oll'e~a'l.l
range'

R,an!]e of

(J;v,era II
Il'le-a,n

--

01ieraH
,rllnge

...............

Mililed !"ioe

plant,

I1fHli!r!i~

Rarnge'ot' Ove'filll planl m'eilll1lS mean

------------~~

11'111:00., IIIAS X

Rikut·g
INJorifil 20 IA~1101,. I IRS, X Omil't:3'9 1IIIR'11102. IIRS X
SeUllQ

~8; 5 1

8.0.113,,4,

9.1-112.3

1'1.1
1 '1.1 'I 1.:2

7A·1,2,7
:EI.4-11 '1 .3

7.9.12 ..5

10..6

9.2··~2.0 10.1- '111. 5

'110.'5·' '1.8 10.2·13.0 11.1·1'2.1

9.0-11 '1.1

10.4 '10.8

auoa

1114,"1.0
11(1,3-12'.5 U:I.!S·12'.4

I FlS X.

CilQw·ru rig
I frl HI4,
KOI1!l

.:'I

11.1

11.5

HU3.x 5 110.2,·1 ~IJSi

UIOJi-l 1.2

Crylhr,oce os 1 '11.00 9,4.1 L6

1

9.9-'1 UI.4

~ll,S

I Rl110S, IiRS,.x Chtok·jye·bi,chal I A,S (Conlr,olli

a
~12

8,,,6·~14.3 7.0·10.3

10, '1-13A3
7.4- ~li.1

~I:i!.~ .

8.7-W3.'5
.. 16-3· 911

tl8,1'3.3 6.5-9.4

12.1 8.1

8.7

,55

and 5.75% lysine, whereas milled rice protein had 21.5 and 3.32%, respectively. Because of the more even distribution of protein in these high-protein rices, the brown rice aminogram is similar to that of milled rice. The glutamic acid and lysine contents of the brown rice protein were 21.1 and 3.53%, respectively. The rice hull has been repo rted to con tain I to 3 percent protein, but the aminogram of its protein is not available in the literature. Therefore, the aminogram of the hulls and brown rice of three local varieties was determined. The hull protein showed higher values for alanine, glycine, lysine, proline, and valine, and lower values for arginine, glutamic acid, isoleucine, methionine, and tyrosine than did the brown rice protein. Since hull protein constitutes only 5 to 8 percent of the protein of rough rice, its effect on the rough rice aminogram is very slight compared with that of brown rice. Rough rice protein, for example, has a lysine content 0_14 percent higher than that of brown rice. Because of the increasing area being planted to IR8, an investigation of the effect of replacing traditional varieties with this improved variety on the protein level of rice grown in farmers' fields was made to test the prevalent belief that yield and protein content are inversely correlated. Analyses of IR8 and traditional varieties grown in the Institute indicated that the high-yielding IR8 has a protein level comparable to that of the traditional varieties. An arrangement has been made with the Statistics Department whereby samples of IR8 and local varieties collected from farmer cooperators in the province of Laguna will be analyzed for protein content. With the increase in the importance of IR8 as a rice variety in Asia, the nutritive value of its protein as compared with that of local varieties should be verified by actual rat feeding experiments. An amino acid analysis indicated that IR8 has the same lysine content as other varieties with the same protein level. Since techniques of protein quality evaluation have improved in recent years, a cooperative project was recently initiated with Dr. R. Bressani of the Institute of Nutrition for Central America and Panama (INCAP), Guatemala City, to study exhaustively the quality of rice protein. Three samples of milled rice were sent to INCAP - Intan, with 5.7 percent protein and two

IR8 samples with 7.3 and 9.7 percent protein, respectively. The moisture content of the samples was 12 percen 1. Three types of PER assays are planned: (1) at the same protein and calorie level as found in the normal diet, (2) using a fixed amount of rice and correcting only for the calorie level, and (3) at levels of protein varying from I to 7 or 8 percent in the diet. Studies of Rice Protein
Soluble proteins

Studies on the various protein fractions of rice were continued. Preliminary work was done on the amylases of mature and germinating rice grain. Total amylases were assayed by observing the reaction of reducing sugars formed with the Somogyi reagent. The bran-polish of mature IR8 grain had a higher total amylase activity than the milled rice fraction. Phosphate buffer (O.IN), pH 7.0) was a better amylase extractant than 5 percent sodium chloride for both milled rice and bran-polish. The amylase activity of the phosphate buffer extract was about four times that of the salt extract for bran-polish. The activity of the milled rice extracts was improved by increasing the salt concentrations up to 3 percent sodium chloride after which it declined. Corresponding increases in salt concentrations in bran extracts lowered the amylase activity. Amylase activity may thus be presumed concentrated in the albumin fraction. In whole-grain and milled rice extracts of mature grain, l3-amylase was readily detected, but a-amylase could not be found by the assay method employed. For a-amylase assays, ~-amylase was inactivated by heating the extracts for 5 minutes at 70 C. In the ~-amylase assays, a-amylase was inactivated by adjusting the pH to 3.4 (ice bath) at least 10 minutes before the assay. Agar-gel electrophoresis and immunoelectrophoretic analyses were applied to the bran-polish extracts. The agar-gel electrophoresis at pH 8.2 of the buffer extract and the subsequent amylase assay of the gel revealed one fast-migrating amylase band. No such band could be detected with the salt extract. Three precipitin lines were obtained with the phosphate buffer extract and only two with the salt extract (Fig. 4). Immune sera were prepared from rabbits injected with a rice protein preparation by Dr. R. Djurtoft of the

56

Cereal Chemistry

Technical IUni!;'llrsi~y of Dc:llIimarlk mn Copellll'lul~~n. A ge~ il'iUra:Uon 0:11. Sephalilex G·loo of the buffer extract showed eli.ilt the bnm1!!rn}dase was heterog~"IleQlIS". Two \?!'CrC ,obtl,li.:ii!ll;~.

bl!i)iI.d pcaJk.s of ,aJmlfla~ acUvi,ty

Jhe' changes, lin the pl)ote:ill i::ra,cttolii$,(hJ~ing g{!nni$111.UQI'Iwere i[:~di.ed also After ~:ne ~cond day or .semlination II. ~,Ii,pi.ddnl:p ruliltlilae gl:u tc:I~i1ian ~ ~obuUIii fr,acUoliis coincided \V,HIl an lnereaae ill Ule level lnoll'protCiin N per g:ntilll ,( ~g.5)., A t:011·

lIIoted. boOth ,Iii, iind O.amyl35es were [Io'ted after ~he foulith day' .of gernIinaliQII.. Dis~: elett[!'opl~ore~is showed two' amylase bands; .in extracts olig~rmina,Ung ri!:c Wllich we~,e liiot d.e[e~tO!bl.e iJII ~hem.a nne Il'a~n .. IFurlhe~ s,ludy' of [,h,~amy.l~sisilll pro3't(lss.
S'l:ilil~. W<I S

or

~e'v.ecl a.lbu min ~,~ :prol<!min or

in

Increases

ILh,c activiti.es of

~:h'lIeQ"(-.d! 'lljIi~h,plul&ptl!UC:
ciM;~ori"d,e.

FIg. 4..

hmntlll1,QI1!lec

INlll~Of,jHnil'

b,rrr~r..

p~.ucrn~~l' brjlnpro ad" ll<r.J.i!I 'ill'ith !:i,~ 'sodi~,m

P,l'OllImiIII

-r------..,,----,----,---"""'iF""-......,
Q"~

Prolamin prcpa.rn.ti,o.llrs were obtained t1'O:m defau,ed brownl andl milled IRS rices wh leh had been previously eX'llCilcted witb water and a. S-pCn~enl
saUsoll1!,t:iol'l. 1().pcrc::ent 91,cohO'.1ooocen t~;lI'I.io,~~:X:I:iI"!l;:li,ed sUgbtly mere ~!JO' ~alnilil li:li,m dlid 60 perce.Du, etl~3:niOl.11111l'C'oovery ofprolomi.1ii WaS, h~gh:~r r~(liln b~{I'wn rice iJl~o'teilii Ulilml r~orJll mn~e(l rice. Previo'llls, stl.ldie-.s ~ilVC ,s'ho\Wilil,
<I

v~
I~O

• L;!ul!illn
",~.~N

t----t- ..... ~_t_-

I~

~I",

-'r---I

Aqueous Ic(bru!lolll~,

tha t btow~1 dc:e p~o~'jll has !l I:l;i,ghe:rOOi1lkfi~ ,of :plu$mnillil t!hu:n mi~led dce pJQIHl:Jn. the diiilyud pro]runil:l recovered \V,as: 0.2 to 0.3 percent of lile
~olal pr'o~iC·illll. TIn~ ·p.repaii'!ltions
l,I,.rcnlv,ery

hyg:l'o·

I,
DIIfts, o!l~ Glili'ffiitiGl~
Fig..
~ G ~

scopic

and C'QJiI'Ia.iIfiCd ,e:.;ubolitydJat'e-s. u1ainly sillCOSC. The: nature of 'thi? ,ca~lbobydIli3fc''fra:edon is under mully. G:I!;Ir:e11i'll
A eomp3~iso.n, of ~]U"ccme thm!s Qf lex~r.a:ct.ing g!ute]11iI from JR_g, mulled ~:i~ 'lWS Y:n,d~'ftaken. l'ile Undn!!.r nttH:in,odillvo,l~d tlhe cxtmot:iolll. of protein WIi~h sooiu,m :kyd(,oltid'e ~O+O] N) ,IJj[ldthc preS d:l'iUU iO,1I of glu tcl:in.pmlanU:fI by addi.l1c.uioll of

S. Ch~JIj]ie~ ,IOQ the pro'lein fract'hons 11:lIIriJ!lg g,e<lT!Il;n~ .. dci:fI 'of lIl,g !lIraill,

~he extr,i!;ct:s. This prcc:il!lit!!:te wa:sW3,siled several times wUh 5 :percclltrodiun1 chl.m:ide and ~'lIe'pro~ 1:I.mi:r!,w~ extrac'tediwiil:h 60perce!l11 ethanol. A mQdinedllLindnc.r procedure invu!,¥eCl the extraction of tha f'loteil11: by pe'r-.;:ot!l!li!!!g 0 ..01 S.N <m,d O.OJ tV sodffiumhrydHlxJodie' S.o]UU.oIWS tll~OUgh .\Ii ,oohil,m:rn (l:[ ~i;G(l'. OsbQ!'J1oe;·$mMhQd. oonii$w)~ of c'.xlrn.(ltil'l.Ba~bm:11Ii~. ~.obuj]ilrl. w:'UJ: prolamilll\l,\i~h W<!te[., 5 percem -~odium chloride.a.l1d 60 ~cili'cen~

willa O.OSN sod;iom hydroxide. 1l1e gl:uteUn preparatlnns 'WCiI"e pudfied by di:s:sobn,ion ~n alk a:li,a.nd prodpUatitolll wit hi :1I,yd.r,ochl.Qrical;;id. Some: of the preparations extracted wl:lecUy from :miUed rice wi:tlil alkali w.itlum'l prior I~xtracti:on of tIII.c other PiJ(i~e~n frae.lioll,S} showed abnooJ<Jllal:ly high I:ys,ine' ccatentvalees 01 6.0 and 6.7 perc,e.n~ ..Tbe' oithc~ ~\I'clPaJ'l'a~i,Ol11l1 lldlys~n.~ ·OOI}~.ciiUS o.f J .4,2 to, 3 :94h ~~oetltt. th.e glulenimJ ,n':pru'led ~y Os'dl~.lrin.e's mediod ha~ i~ ]l!igher N ceatent thrlrlil ~litJo:~ of ~.h0 o'th~r prCp:3'!1!:I.ioUii.

~lb;mo'l, rcspeC'livc~~~bdon~ eN.[~~tU~8~he:glnJb;:l.in •

O~nilJQlemist'r_l'

51

The intrinsic viscosity of the glutelin preparations ranged from 12 to 20 ml/g in O.IN sodium hydroxide at 25 C. Ultracentrifugation in an alkaline solution revealed only the 1.3 to 1.6 Speak for glutelins extracted with 0.0 15N alkali. Those extracted with 0.05 or O.IN sodium hydroxide showed two peaks with S20 ,w values of 1.4 to 2.3 and 4 to 6, respective Iy. Preliminary lightscattering data indicated that one of the glutelin preparations had a molecular weight of 3 x 106.

Preliminary

experiments

were made on three

Protein metabolism

In the Institute's intensive breeding program, lines are being produced which are essentially isogenic except for a single property, such as protein conten t, amy lose con ten t, resistan ce to rice blast disease, or resistance to stem borer attack. Such plant materials are ideal for studying the biochemical basis for the differences among these properties in plants, since the normal environmental and varietal differences have been minimized. The first project, the study of protein metabolism, was initiated this year since lines differing in protein content will shortly become available from the Varietal Improvement Department. The higher pro tein con ten t of some vane ties possibly resul ts from a higher rate of synthesis and/or a slower rate of breakdown of protein during grain development. Protein synthesis, in turn, is affected by a number of factors, including the level of enzymes along the synthetic pathway, the availability of amino acids, and the level of ribonucleic acid.

varieties, Peta, IR8, and A2 torzs with a low, intermediate, and high protein level, respectively. Planting dates were adjusted so varieties would flower simultaneously. The plants were grown in pots in concrete beds under continuous flooding. The overall protein-synthesizing and hydrolytic activities were assayed during grain development at 4·day intervals using intact and homogenized brown rice. Samples were kept in an ice bath during the preparation. The capacity to synthesize protein was determined by measuring the amount of radioactivity incorporated into the grain protein after the incubation of intact, hull-free grains with a complete mixture of amino acids containing leucine-D-!" C. Proteolytic activity was estimated by the rate at which a crude homogenate of brown rice could break down added protein. Assuming that differences in the ability to synthesize protein are found, the synthesis of ribonucleic acid will be assayed by allowing brown rice to incorporate adenosine triphosphate-' 4 C from a mixture containing the three other nucleotides - guanosine, cytidine, and uridine triphosphates. Ribonuclease was estimated on the basis of the ability of the homogenates to break down added ribonucleic acid. It was found that the ribonuclease activity in the rice grain was much lower than in corn. Proteolytic activity also seemed to be low. These preliminary experiments provided the bases for the selection of a standardized method of assay which will later be applied on the isogenic lines differing in protein level.

58

Cereal Chemistry

y,eaJ'. S'lx pedigree

't!he breedin,g progra,n lWS cotlsid,erabJy exp.IJ~~dl!dd'tlrl;~g ,the mJtseries ~vU}! a .to.laJ of 41,.571 r.ows-

selec,r,ed from 324 iliffe,rel.l:tcro:sses were groWlJ. Alotal:of 5:5 Fl popuitJlions were planted andover 4.0()O plant· £'elec#on!; were made /110111.' .tlf!e~nf(»'gro'w~~ng ir:r {J:edigre{~ row:s·. So't!JrJ:ofthe i'nt[J'O¥/:(JtJ Z' H,ew crosses W'€ir'\e betw(!(!I'J !RB . (ll'ul ;six }f'igh~porlefn v(Wi(tlieSi.Se~~,eral.F pedlg,r'€,~rows 3
/10'.'m these crosseswere of NNJiSonab.ly :good ph:m,r type .(jl~d had as high a prm'eil! coutetn CIS12 IJ/!,r,cel'J:t. In ,the sat.lte

e~~p:enm€nt l.he. pl1otei11 confirm/of tl~(!/.R'B cl~(!ck rows paried Irm",?::9 to 9.1 pe11celJ'i.
Obserm,rtonal
a1.J:d ~ii!p~tr:a,led",ielei

.r:lurltJg.tlte year. Selle.mllit.teSc ,(Jlgoed pl(m t .type y.ielded

'hleUas IR8a.nd
dis€(J~s(1rui
i"ll

t.rial:s wer,e .growu twice

.(1;S

(;lddi'lionshm~e·d resistam::€

to ~e'lterll,l
.milling

i~r;sect:.5 ami pmse~sedg'~'ins()lirtl'prov(f1(:J

aru:l ,000 ki"~ quaJ'i.ly.. These p,rQmisbtg .ii1'!t?S,are bei1~:g tested i~ ITu; ri;cegro'Wbur c()wUl'ie$ of AI si,Q(11,1(1 SomhA mer/ca. und it is hoped that one 0' two ,(Jflh(!.~~ will be jound su#abJe for release as ne \VI!(lri'rNies •. Pur~seed progF(UI'I,s!or $ftl!'er(J1 p'f"O,'nisi11g' s,e.(e·ctions were s.tU1'.I,{!,d. b~1968oll·er 12J)()() samples of seeds of J,Mli't[ue b,.,eedillg lil~eJ~vere sent to 1,83 !j:ndfll.id~'U(! Q~d (},rgan:iza t.icm~'ill' 79 ,c:oluu'~ies\, .11. large l.i'umbe·rof Mrlel.ies al:l:a l,lu~s wer,e sent 1'0 Ml'iO,usctJoptU'tdir!g' COiUir! t,ies tt» de.t,ermtne tt~et,acklp lQ,btlUy, tes.tsl:cmce tOIl'aJridu.s: diseases. rflactiQnlG low nm~p€.ml'Urf!s Qfj(Jtor perjo17.nance umlerccmdltior!s of deep' water. In ad'dUlo'1.i~,. !>pecia.l'lUf:s.e.r,i.esof bj',eedll~g itnes wen' sent to I:he coo,P.e:r:atil'l\gcout.ltflf:1:s.!().r pe,ld lrtstsfor n::siJtanc,e to Me t;(1t:fall:eafb.Ugh t, blas.~, ,and .the umgro :I'irus. Getuutc studies to aetermltJ:e ,rlrea#ell,(;: .mlatiomrklps of )''a"IOil:lsdwarfs ,rma $,en'!i,dwarfs ,and toasce.rt,ain tt! e inherita'M:·e pauettl ofplwl'OtJer:lod st:l'lsitt~'itv lv~r~ ,oonlin!:t,ecJ. Cyto.l()gi'C~'1 il.II~(!stig,aliorJSof the causes 'of ~term Iy iJ1 the sterile indica ;.-.;: i'u;Ucacro.'5.s~$ werec.o.n linued. (litO ,'.lewS:(r.ulies an

the ~,t,erile indic~ x Im/'f,t J':,/)!f"i(/S were initiJj,t,ed.

59

Varietal Testing and Improvement

World collection

The number of cultivated varieties of Oryza sativa maintained in the Institute world collection now totals 11,057 accessions, of which 226 were received during the year. Of this total, 652 accessions were grown during the year and morphologicagronomic data were taken on them in the field and in the laboratory. New accessions are grown in at least one wet and one dry season for proper description and classification and are regrown from time to time for maintaining viable seeds. A varietal catalogue which describes 8,628 accessions is nearing completion. Viability is maintained by placing a 250-gram seed sample of each accession in a cold room maintained at 1 to 4 C. The moisture content of stored rice is gradually reduced to below 7 percent during storage by placing samples in sealed glass bottles (2-gallon capacity, with 12-cm diameter screwcaps) containing silica gel. Seed samples of 12 accessions are placed in each bottle along with 600 grams of activated silica geL The silica gel is replaced every 6 months with reactivated silica gel. The moisture content of the rough rice at the start of the storage period usually varies from 11 to 13 percent, or the approximate moisture content reached in the air-conditioned seed room which is maintained at about 25 C. After several changes of silica gel, the moisture content is reduced to about 6 percent. An experiment to determine the viability of stored materials over a period of years was started in April 1963. Seeds of the varieties Peta, Siam 29 and Chianan 8 were placed in air-sealed metal cans and stored in the cold storage room housing the world collection. Each can contained 1,500 grams of rough rice and 500 grams of silica gel, The cans were opened every 6 months and a small seed sample withdrawn for determining the percentage of germination. Reactivated silica gel was added each time the cans were opened. In 1967, the me tal cans were replaced by 2-gallon glass bottles with metal screwcaps similar to those used for storing the world collection. At present about 1 kilogram of rough rice remains in each bottle. The amount of silica gel placed in each bottle is 600 grams. At the start of the experiment, the moisture

content of the rough rice ranged from J 2.29 to 13.83 percent. In July 1968, the moisture content ranged from 3.08 to 6.33 percent and the percentage of germination from 83.8 for Chianan 8 to 95.4 for Siam 29. The germination percentage of Peta was 91.2. A total of 500 seeds were used to determine the germination percentage of each sample. During the year, over 5,000 seed samples of accessions from the world collection were sent to 147 organizations located in 60 countries.
Breeding program

The breeding program was expanded considerably, with larger numbers of F2 populations and pedigree rows being grown during the year than in any previous year. Similarly, the number of breeding lines grown in the observational and replicated yield trials was also larger than in previous years. The seed purification and multiplication programs were expanded to involve many new selections. The main breeding objectives as ou tlined in the previous reports were continued. As some of the insects such as the brown planthoppers and the green leafhoppers act as vectors of rice viruses besides causing direct damage to the rice plant, the problem of breeding for resistance to these insects received considerable emphasis, The breeding for high protein content is another breeding objective which received attention during the year. The crosses of six high-protein varieties with IR8 were screened in early segregating generations and the selection for high protein content appeared to be effective. F2 populations, A total of 55 F2 populations of new crosses were grown during the year. From these populations over 4,000 plant selections were made for growing in pedigree rows. Over 700 F2 backcross populations from 35 different backcross combinations were grown and over 4,600 plant selections made for growing in pedigree rows. Some of the important new crosses and backcrosses were between IR8 and six high-protein varieties, and between Peta/3 x Taichung (Native) I and Thailand varieties such as Leuang Hawm, Gam Pai, Khao Khian, Khao Sew, Khao Poodhat, Nahng Suwon, Khao Med Lek, Leuang Pratahn and Khao Dawk Mali. IR8 x Intan is another promising cross of which F2 bulk populations and F 3 pedigree rows were grown.

60

Varietal Improvement

P'ed$gret. RU,rkties. peditv,ee liIU~iPe,rjes Wl:r,~WWTiI iII! November and Delee:mber ~967 <lJ!ldm Fc'1DlrYU)I.
MilY, lhdy :a~d A.u!lJlls~.. rn.'968" 'Fhe f.luro~~ of' ])eldi·

'1'Q.Me 1. le~din.i!! ,~mSllell glOW.ll i.1II lledigJee IRRW. N~mbe( - ~967 mill '\U.~J1 l ~6a-

:nl,hfse:ri.'es:"

:gre_erow~powl'! durirng:th~~ fC:riod Ito~:wed4[ ,57'7'. or tnis ~1LI:mb~'[.'2iO\8Q H'WS c",ruine>~ ofpare:nt;i]
checkYll~ielills .$pa.~\di eve'ry 20th ~OlW. 'rb~pr:m~~ lines grQWUi!. Ur1ngthe yeaII' w~~e d ~I!! ctc,~ Iroom 32~, diIfi:l~fl'tCfOsseS ffla.de atmt;l mnrs;tUy ee, The ~"!I'ge Ilu:mk',u of CIlN£e5 in\l'olvedl p,~~ ::;om!~idea :~,[Q,the .e.enetic diYenity olf the: modUu te's b.n:;~diQgpjog:!'!1mJ. U iseS&!!:nUaI, U:UII~II. wide l'.:lllge ofgc]1~ti;(l diversity k dcvelo~ed witrunthe im~rovedpla'nl· type Mlltnria]. Tbls di.!/CI~ily illCl]l1Idl3~ wjde dMfeJ" enees if.! 1Hi.s~(l,liH~e to di~~s!!'f:idin~c:;t~.M~i!f 'cha~aclier.i~r!k-:s saeh as ~IO~Bhml.es£ ,adr.ue of '~.n,e~n:ce! d~gree of refOiUil1l:1ce cold! ex:tJiemcel.y te wjde YiJlI'iJariOliliS In grain ~e1li!'!d ~iI:<l!P~, m_illmiG aI'Id .Mld ,eoo:k:iUllgpmpc nlrM. Ei;gh1.y @f mle mO!le p:rDmi~m.g C:I'O!5WS~C'W11! in pewgI>e'e WM are USl~(:I in T!l!bl~ I. ~O$$eS r~p('~:!1i,~, 'O¥t;)r ,65pefceml @r Wlet.ota] Immbe~ .of

No. of ped ig!!"ee I1OWsgrown

IR915,
~fl26Q

P!Q'La!2.ill TfN ~1 Pet;a!3 x TIN ~1

2,4

36
51

iR400
~.A482

~tal4

)Ii

'liN~1

P,erw5 X l'(N) 1

~,A564 11,1'1,8'58 Total IAH)913 u:l5;n ~R578 ~,FUK~O

PetalS", TrN ~1 P,etafl' xl(N) 1

45 7ll: 467
70]

mao K lntan 1122 IIIRS.x SigiPdis 113:2 1m S x [Si'9illd ls ~ 'nl~ W]2,411 I me.iN: 1~IMlC;VA xHleo·IIZi!'~ UR 12·1'7.8) 5118 ~[!20
11'1:8 x [I ~4·253 !'i IB589A4· 118· 112 xl(N)

n~

SliJect[on$. gJ~w;n.
"FheP~tll ~ TaicblilliliS ,(Nati.'i'e) ~ rDil.C ~(i'r~· ~Q hta mi:!ideuf' Ql1Ily 70,~ ~o,ws but aside f~om ~.c:

01

401

IRIS3

PeHln x T<rlchum,g. (Nat&ve) lcombiR<!ijo:n, io· I~nsi\l'c seleetten hall bee IJI cl!j~colnum:!l:d. M;iI!!il.y ,axue'm!~my p.rom[siil'liS Ili:n!~swere ob~a:incd froml d'liis serjcs or bil:ckc~05~"~.Tile'), ~lilI18(l m~tliiril¥ fr,o:m, Ln. 10 day~ e3!!'I.i.e~ 15 O~' roO're days larer ~m ~o .UU!,
Most of 'lhe~l:ine~,i!ire' n~~t!!~U~Ql~J!!lJo~!'!d(lire mo~e' r1:~~$t:mt to .~:ssy $~Un1thaJ~, nUl. 1"he:y }L~'VC' ,e,X!ceU!:li~U ~dml ~ype ill'ld~:I;~r'lity :5I!.em~' blrniiilfe: no~. ~y 'retist;mt U) th~ rt.la.ctenta}diseases. I"HH~:nilly.,. Un~or We Peitax. '[aicl:lI:m:g (N,at~e) I backeresses ]lilive b~n focund wm&:h dOl nat ,~ow

IR8!2 x rli~4 •.i!53)1:; (e58~A41·,1:s;.~/2 x T(N)1 }]72 I R400 II: IU 1'18,.)1. I fl.4-:2:53,i
II

j,IB5.89A,4· UH

12: I(
244 7'U

1"1~NI '1~I] Tota~ m7SS

~A759 iIlA.879

H~'8'2
I~R643 IIIR:l419

m~whi'te
$~

be!])! cl'l:iLvac:tc.isti,c fthe gcainw~Lich ~s IIRSOO o p:rominerl't 01'1 lIS gruilns•.,some of these cleawilotal grlli~ 1ili1:l0S,are, b~iI1g,rested e:xtenSi~e:ly :ti,~ the .1JiI· stituteood elsewhere. The benet rehl li;. T,.u~buWl;g (N'aU'l;lie) .~ b!tCkooiSli linlls arem:msid:erI.lOl to be octIIR626
s~\'I1IJ:d~;IiI.B, parerrt as :nultell'iat
Jrn.~.!ll!j .~!l';):rtc'~y clo~lY!Ml,l~t(ldito P~ta, was IR665 c[',ossel!li 'Wjih IR.8. F ~ pewgreeLmes Hom, thi:s C~Q$$ were gJ1ollVllli'll tbot: Au:~ s~. liIursery <illdlJ!ldUlctcd IR151 many good. ~cmi'grain ~l)'pu whidh ~o :no~.l~aJv~

mOJ:!!!!: Daw,i11 275, III 8. x (P~taJl.3 :J( Dawn,1 R 286 ~R 812 11;; (f'eita<Ja.1\ Dawn.1 ~03, U'ISx: [ilal!.l'll'rI x n~',IH:M7 I~ A !!; B·25 I! Otl!lwni)4BD II A 8:f2 x I~;S 8·25, .xDawn) 668 ~ mRwa: -It (,e~8·25 x '~Iawii'd220

a un

:.1379

U:t,B x j PetalS IBe,1111~ Paitllal,

)!

I FIB x jPetOJ15 x Bel,r,e iPatif'iIa~ I R 8/2 )I~ (Pet1il/5

e,eUe P.atmall

>l

wh.iitc beU)'. Som~ cflh~S(J! 1J!l~ U~~.d ~Q i5ho,w klWiI!'I amylo~ C(lillb:n,~ (hillll'l l!i.eP·'eL3 hybr,id~. They offer mu:t:hp~o!!'!lise as ~\llr~lJ'U r.ruttJ!riatland 3ll ~bi!e:oomrnerc iill varieti-es.'flileir only S(lriIOU$

~Aal3x (Pet:al5 x
B~II~ P.a~ii'laJ

585
2:394

Tota~

IR no.

Cross

No. of pedigree rows grown

IR No.

Cross

No. of pedigree rows grown

IR533 IR661 IR594 IR593 IR755 IRll3 IR874 IR878

(CP-5LO!2 x Siqadis) x {Peta!3 x T(N)1] I R8 x (CP-SLO x Sigadis) IR8 x (CP·SLO x BPI·76) I R8 x (CP·SLO x Nahng Man 5-4 I R8 x (Cp·SLO(2 x Nahng Man S·4 I R8 x (Cp·SLO x Nahng Man S·4) IR8/2 x {CP·SLO!2 x Nahng Man 5-4 I R8/2 x (Cp·SLO/2 x Nahng Man S-4

IR748 141 506 82 Total 611 IR627 IR752 IR881 IR793 Total IR1154 IR666 IR568 IR667 IR781 IR934 IR583 IR800 IR672 I.R450 IR744 Total IRI59 IR520 IR 1001 Total IR579 IR532 IR580 IR747 Total 519 811 IR52B IRB77

I R8 x (Dawn x Pankhari 203) 174 I R8/2 x (Dawn x Pankhari 203) 254 3831 IR8 x Wagwag I R8!2 x Wagwag IR8!3 x Wagwag Wagwag!2 x I R 8 251 319 154 143 867

}B3B
2178

I R B/2 x Zenith

1127

Total !R754 IR835 IR841 IRl113 I R810 iR838 I R 1109 IR844 I R 1108 Total IR848 IR789 IR887 IR933 Total IR574 IR822 IR932 tR854 IR825 I RB x (CP 231 x Dimal I R8/2 x Pankhari 203 IR8/3 x Pankhari 203 (IR8 x Pankhari 203) x [Peta/3 x T(N)l] (IR8 x Pankhari 203) x [Peta/6 x T(N)1] I R262 x (Cp-SLO x Gam Pail I R8 x Muey Nahng 82M I R 8/2 x [Nahng Man S-4 x T(N)1] IR262 x Leuang Hawm I R262 x Khao Dawk Mali

I R8 x Yukara 273 Yukara x T(N) 1 270 IR8 x [Yukara x T(N)1] 692 I R8/2 x [Yukara x T(N)1[ I RBI3 x [Yukara x T(N)1] IR8xFF33 Ch. 242/2 x FF 36 (Peta x PI 215936 x eh. 242 Ch. 242/2 x Peta Ch. 242/2 x Pankhari 203 1477 613 168 143 51 87 103 3877 Basmati 370 x T (N) 1 Basmati 370/2 x T(N)1 I R8 x T-3 Basmati 126 75 68 269 I R8 x Tadukan (Peta/3 x T(N) 1 J x TKM·6 IR8 x TKM·6 TKM·612 x T(N)l 463 1151 527 138 2279 [Peta/3xT(N)1] ADT-27 x 223

172 51 338

I R 262/2 x Khao 126 Dawk Mali 56 I R 8 x Khao Dawk Mali IR262 x Khao Tah Haeng 247 17 I R 262/2 x Khao Tah 23 Haeng 17 342 I R262 x Puang Nahk 16 11 IR262/2 x Puang Nahk16 1366

336 309

I R 8/2 x Muey Nahng 62M 236 I R 8/2 x Muey Nahng 62M 881 294 1942 131

62

Varietal Improvement

No. of pedig:ree
r'O'IllI"S'gHlW11i'I

1:01)

H;I~(105 ~R586, IIU191

1

' HUI!!~(FB·2~ )( T., 172)

UU! x UlJiI':7 li/:2 x T. 17.2,1'333:

1160

m:s x

I~BPI'·76x, 1. t76)!121

TQMII

60~

11~!!l:89
1~i'M74

:lfl47B
Total ~.R436, 11'1441

Su~;aflil!andl~ i(lfl\JlJ 1 x :U5 OP..SLO x Sulka11l,an,di199

seL~c:ijoos f~om. Sasadis x T:(.:ichl!l!llg (N:(l!~:i¥e) 1 rba.ck:t.roiSs (Ut-30S), IconWllUe 10 .owpromi.se." Some o:fl'he !bette. lines. ;ure·Wghe.f lille.r.mg ad m:o.u:cr .iJi!. h~~fjbtthM IRS" n~Ya!re Olils(lrno~ ~~ist:am~ t>o ~!e bllcted:aru l~.a!f diseasel$ amd blast, 'J'tJ.e:se hes ~3~ y,ietde:c!I weUin vaJriety and f,e~li· lizer y.i.eld hial~. 'Ylh:il)1.l ~_,e)' may n'o~ de;\le~Dp.m~ cOllTIllcroiaii vati~ties" t'.hie<y p:I'mdd!e anolher 'va[lI;· .aib\lepil~[Ibil:l;oulice. lIR8 ;Ind ~ Ple~~/4 x l'aich!l,][!g (N31tiV1l!) 1 lillTle we~ecmssed. withlhe Flprnilillt ofa J~:~:!:w~~n l1Ri4·2S3 (M-iOS xl!lee·~o·woo-gen)' and an adVa1llilC!ld: gel1ic~Si:ti.om.sd~e~iOJil. f:rom IJ;5S9 A418·l12 ).;.'faichWl,g (Nati.ve) 1. Ex.ceUemIt.I:ilnlesoomJ~ , bioif:lS lR8pllU'it type with g~Qd ~!~t.!!g ,gtili!'!$we,1'i': !ob~:i!iflcd.n wa~ anl~i.pated il:a:~. biillC~iIl!rj3i. .Ie,a:f b:~ghtresJ!$t8m1C~ w,Q'tl~dI be Qtm:lIiJ1!lild as.1I;589'A~·18J posse:ssedresis!tDce. Howeve~,lhe 1I,:;:89A4-18-] f2
x: TaiC:~IJ!ilg {Nati\l'c)1 lse:]ecti,onl used (lid not ca.m)' re$~ti.lnceto, ,the ,tWiw~. H·~ ~,ppe~lJ~to have i! 05 high .~IWtL'lI. f ~le$ls:rMoe I~O IDe broWflpL!l:JilIw,oppc~ o (JV,ilfJpilnatac li:lpns) :.md l!he lines selected f't'Qm th~~~ijs~SI w~tffil. UUI l!!!ldi P1!;~lIl/4ix. Tai:C~l!I,flg (N.~:t'ive) ~. menlioa:Jedabove wliUJ 'be le~lied fO:l"

CP"S!lO )( P'il1i91aew56 (;P.;SIlO x Leo Mue Nahng

nlNIl ~~ ." teb M'~!e ,~lahn9

24 60
63 282' 429

IR4::J9!
1A:442

[Peta{.:2'/I: T(N l11]

;II;

Ll!lb

:resist<lnceto the laseet, Theva:riet.y[)l'!w~ h!l!5 I!itCe~used extenS~lji'~ly as a Mil5IB!e:sistomlpa~ent. Ov,er 2.300 IO'WS of seleet1)QIi!$.

from Cf'l)slle:S .in¥ohiiflgD(lJW1iI

\W:re~:I'OWIIJ.mfl

Ildd:l:liolll to

blast lesist:mcc,wcseLmes

mow

f'lQ-

mi~iflg [')bnt Uai~s ~cb as,~()t1wse!:l~:ooclilce,

~IOlllgh

le!l¥,es, ila:Jdl~ofl'In~bescentpll~:nt

!I:larls.M!lfI'l( Hne~

,of gxceUe:n1( grai~ :sh~poe ~!d 3:W(!;ili[!lII1,o~ a:SW'erul. as ]OWil!!f :a:Il1ylow content ha.ve been fou:nd ilI1t1:1e~
C:I~!i,

fMm,Petai/J x t!JJn~o, ,arlil~ d.'HlwbaCk:i:S~i'!i:t lID;:];: Offes_.isl3l1loeto Ihe bi:lic~'[ial gra£!sy5W:lllt. I!!nd !l!1~,wi!halil·81iB·25 x Dawn! Ilae, 111: bQtl!i I>om:bilullilion~, one b.u::li:.c:l'l;}S$ L'O [R8~sr dii~~:s, AmI,o~herseJ~c'l~on "!V,hiclil mO''!!iIe:d !\In el((;e~le~t !1nl.!~, Mull' f'loroisi!'lg 'i)1:!IS:t.f.~:d:s:tant s.elieiCtiOilil~ .g!I"!1iintYf~ Qf~owe-J:~Yruose CiQ~l~:~~ Hum :IRS 'was we~ema:de and .aJwta1 Q\!1cr 1.700 line~were ill dW311'f line f~om the (;~(}S$ IRS :It (Mo~ ,Cbillm ,grocWT! af!ds~l!Iid!i.ed. Van8 A x I,gco.tllfl). ,["me .Mons: Clhjm y~ A.x An ea:dy malurmghtl/S X llellJe PaU'!~line: W.11!S I,"&eo.~epa~er!t U!Ile (nU 24 78·2~3) W8sms:ist:an;t oro~d. withrnR8f;oUOMldj by oae and two back· ~o gJ'ilssy stulilt.tite~.aJl.'!iI$ m.d not shQIW wl~rute eresses kI niB. A '~Q~:3'lI of 2,394 imr:llCS 'WJe~~wwn g belly, md tile anTIiyllo!l6 oon~ll~ wa.!l24 b) 26pe,[· from. these conibmlau.<'lIl5.. il5lH'lym<l!llurity w~IQ~t Ci'Jl:lt oi)mp~[Fed with 30 pe:!'C~~t o.rmo~1l1 for lR8. inm;o~il: cases b11l.t\axccllc~:t, .IIOllIS, rull'J\de~iGiI'3mtype'.~ s of good p.lalH:Hype we~eObhi~.ed, tM~lm~sha,v~ MililWY good iPbnt·~e~[~!!c(!J:n]~in~l:lig graMY itunt pi'!~rli)rmed vs.:y W'i!U ~r:I yiehl! trialis. M!iO.they ~~sta:li1Joe.)iar!te~ldibJte :~~y.lose C:~I~~!Th~.~alld absenoe afwh:iw tllilly h~e ~ell1. ~illecll~d! from appear' ~;ohlllveto!l,l;gh~r ]~.e$ :aI!IdmQ~er\l:Siistllilil~
II ]IDIilC

lR8 wa:s ,cfo~d 'with

Daw:n. which

Wl.!S resisl<l!lllt

~~ b~!li$tt

or

ills ((ross,

to' :gra£S¥ :sttl~'U,Ih::IIn.

nt8.

The selection ep231 x S1.O-17 (Ace. No. 6993) has been used in many crosses. It has a relatively Short height which is easily recovered from its hybrids. CP-SLO has a tough leaf which has been transmitted to many hybrid lines. As demonstrated by IRI27-80-1-10 (CP-SLO x 8igadis), clear or translucent grains have been found in many lines from crosses involving CP-81.O. The cross IR661 [IR8 x (CP-81.O x Sigadis)] appears to be an outstanding combination. Lines that are earlier maturing than IRS and have excellent plant type and grain characteristics are being found. Aside from resistance to bacterial leaf blight, tungro, blast, and the brown planthopper, the better IR661 lines combine most of the improved traits mentioned in this and previous annual reports. Another outstanding combination is IRS x (CP-S1.O x Nahng Man 8-4). It appears that some of these lines possess resistance to grassy stunt derived from CP·SLO. Many lines from this cross have excellent grains with clear texture. Some lines are resistan t to blast under Los Banos conditions. Several Thailand varieties have combined well with IRS and Peta/3 x Taichung (Native) I lines. They include Nahng Mon 8-4, Leuang Hawm, Khao Dawk Mali, Khao Tah Haeng 17, Puang Nahk 16, Gam Pai and Muey Nahng 62M. Khao Dawk Mali, a variety rated as having excellent cooking properties in Thailand, has a rather low amylose content and a relatively high gelatinization temperature. These traits have been recovered in many of the hybrid lines grown. Leuang Hawm, an aromatic Thailand variety, is being used to transfer aroma to improved plant-type lines. Muey Nahng 62M and Gam Pai were being used to combine the waxy gene with improved plant type. lines 14 to 20 days earlier maturing than IR8 have been selected from the Muey Nahng 62M cross. Muey Nahng 62M has been reported to be resistant to gall midge in Thailand. Variety Pankhari 203 from India, which is highly resistant to tungro, was used as a parent in several crosses. Over 4,000 selections from Pankhari 203 crosses were grown. Pankhari 203 did not com bine well with IR8 (sterility in F I) bu t after one backcross to IR8, some good plan t types were obtained. Only occasional lines from the IR8/2

hari 203 backcross showed the low tungro infection of Pankhari 203. These lines will have to be tested further to substantiate this reaction to tungro. Both IR8 and Pankhari 201 are susceptible to grassy stunt and, in most cases, plant types of the hybrid lines were inferior to IR8. Except for the possible tungro-resistant lines, most of the Pankhari 203 lines will be discarded as they do not show the promising plant types being obtained from several other crosses. The variety Wagwag crossed with IR8 gave many promising plant-type lines resembling Wagwag in grain size, shape, and appearance. Further testing is needed to evaluate the disease resistance of these lines. However, they Should make good parent material. Some photoperiod-sensitive lines were saved from the Wagwag crosses. Both IR8 and Wagwag have high amylose content. Zenith continues to be a promising source of resistance to bacterial leaf blight and blast. F 3 lines of the cross IR8/2 x Zenith (IR 1154) we re sent to Indonesia, Ceylon, Malaysia, Thailand, and East Pakistan to determine their reaction to bacterial leaf bligh t as well as to low temperatures. The results have not yet been received. Crosses between japonica varieties and IR8 may offer much promise in the development of improved varieties for sub- tropical and temperate regions. If the vigor of IR8 can be combined with the cold resistance, tough leaves, and other desirable traits of japonica varieties from temperate regions, sturdier strawed varieties capable of utilizing higher rates of nitrogen could result. The F I generation of the cross Yukara (japonica) x Taichung (Native) 1 was crossed with IRS, and many of the resulting F 1 plant were backcrossed to IR8.

Pank-

A large number of lines from the Yukara crosses have been sent to East Pakistan, Indonesia, Ceylon, and Korea for determining cold resistance. Stem borer-resistant, improved plant-type lines were developed by crossing IR8 and Petal3 x Taichung (Native) 1 with TKM-6. The [Peta/3 x Taichung (Native) x TKM-61ines (IR532) were not as sturdy strawed as IR8. IR8 x TKM-6 lines (lRS80) appeared to be sturdier but in some cases they had inferior grain types (opaque endosperm). It is possible that lines suitable for release as a commercial variety will be developed from the

11

64

Varietal lmprovement

Table 2. Yield data of lines grown

in observational

yield trials.

No. of lines grown I Rna. Cross Dry season Wet season

Yield range (kg/ha) Dry season Highest Lowest Mean Wet season Highest Lowest Mean

IR95 IR262 IR400 IR482 IR564 IR658 IR305 IR442 I R506 IR644 IR533 IR594 IR579 IR157 IR481 IR1BO IR480 IR665 IR626 IR751 IR875 IR577 IR578 IR580 IR586 IR589 IR593 IRB61 IR532 IR874 IR8

Peta/2 x Peta/3 x Peta/4 x PetalS x Peta/6 x Peta17 x

T(N) T(N) T(N)l T(N) T(N) T(N)l

1 1 1 1

18 25 11 9 7 4 7 47 7 11

8 10 14 12 9

8486 8280 9457 8388 7908 7340 8416

19

Sigadis/2 x T(N) 1 (Peta/2 x TN 1) x Leb Mue Nahng IR8 x [B589A4"18·1/2 x T(N)1] IR8/2 x [B589A418·1/2 x T(Nj1]

6588 6604 7288 6796 6426 7181 7833 5653 6442 7938 5864 6200 6958 6698 6680 5720 5698 4928

7537 7442 8373 7592 7167 7261 8125 7048 7733 8823 7059 6822 7425 8126 7337 6916 6413 5171

6764 5873 6151 5640 6061 6815

4319 4997 3541 1912 4927 2441

5542 5435 4846 3776 5494 4628

41

8442 9024 9707 8254

5977

1911

3944

[(CP·S LO )/2 x Sogad is] x [Peta/3 x T (N) 1] 7 I R8 x (CP·SLO x BPI·76) 6 IR8 x Tadukan 6 L. Yai 34 x T(N)1 16 L. Yai 34/2 x T(N)l 7 Nahng Man S·4 x T{N)l 38 Nahng Man SAI2 x T(N)l 8 I R8 x (Peta/5x Belle Patna) 5 I R8 x (Peta/5 x Belle Patna) I R 8/2 x (Peta/5 x Belle Patna) 3 I R 8/3 x (PetalS x Belle

5 9

7444 7891 9554 7994 8112 7128 5412

4807 5770

3553 4325

4180 5048

7 3 4 6 11 6 7 22 18 11 17 5 13 19 6 14

5897 5341 6202 6122

3862 5043 3602 4377 3167 2124 4556 3770 4294 3142 4137 4500 3553 3382 3022 5253

4880 5192 4902 5250 4359 3324 5297 5138 5322 4196 4968 5269 4832 4644 4540 5821

5612

5002

5307

5550 4524 6037 6505 6349 5249 5798 6038 6111 5906 6057 6389

Patrial
I R8 x Sigadis IR8 x [Sigadis x T(N)llIR8 x TKM-6 I R8 x (BPI-76/2 x T. 172) IR8x (H·l05x Dgwg) I R8 x (CP·SLO x BPI·76) I R8 x (CP·SLO x Sigadis) [Peta/3 x T(N)l] x TKM-6 IR 8/2 x (CP·SLO/2 x Nahng Man S·4) Peta x Dee-qeo-woo-qen 11

8618

6799

7709

TKM-6 crosses. Sukanan di, a bulu variety from Indonesia, was crossed with Taichung (Native) 1 and CP-SLD. A series of rather promising dwarf lines possessing many of the traits of bulu varieties have been developed and are being tested in several countries. Bulu varieties possess tough leaves, are coldresistant, and do not shatter. The IR442 lines from the cross [Peta/2 x Taichung (Native) 1] x Leb Mue Nahng, a deep-water variety from Thailand, continue to show promise. The floating trait of Leb Mue Nahng has been successfully combined with the dwarf gene in this cross. Some of the CP-SLO x Leb Mue Nahng lines (IR441) combine the floating trait with the tough leaf of CP-SLD. Lines from the CrOSSChianung 242/2 x Leb Mue Nahng may possess both the tough leaf and the floating trait. If this be the case they will be crossed with the better IR442 lines to further improve the strains and make them suitable for growing in areas with one-half to one meter of flood water. Observational yield trial Totals of 569 and 621 promising selections were grown in non-replicated, observational 6-row, 5-meter-long yield trial plots in the dry and wet seasons, respectively. Most of the selections were hybrid lines taken from pedigree rows. Data were obtained on yield, agronomic traits, disease resistance, and the milling and cooking quality of their grains. Some of the important crosses and the number of lines from each cross which were grown in the observational yield trial are listed in Table 2. The table also shows the highest and the lowest yielding lines of each cross as well as the average yield of, all the lines from each cross. In the dry season 67 entries yielded over 8,0()() kg/ha. IR8 was in 11 of these entries and averaged 7,709 kg/ha, In the wet season 95 entries yielded over 5,5()() kg/ha. IR8 was in 14 of these en tries and averaged 5,821 kg/ha. All of the high-yielding lines were sernidwarf and resembled IR8 in plant type. Some of them had excellent grains and a few showed high levels of resistance to several diseases. In addition to the two main observational yield trials grown during the year, 392, 145, and 533 selections were grown in unreplicated plots seeded

in November 1967 and February and May, 1968, respec tively. The incidence of grassy stun t in these plantings was so high and the general growing conditions so poor that reliable yield data were not ob tained . Therefore, these plan tings will be discontinued in 1969. Replicated yield trials A dry-season yield trial of 189 entries and a wetseason trial of 162 en tries were grown in replicated 6-row plots 5 meters long. A 30 x 15 em spacing with a single plant per hill was used in both experiments. The same land area was used for the two experiments which were transplanted on January 18 and July 26, using 20-day-old seedlings. A total of 120 kg/ha nitrogen was applied to the dryseason experiment and 90 kg/ha to the wet-season experiment. Basal nitrogen applications of 90 and 70 kg/ha were made on the dry- and wet-season crops. Nitrogen topdressings were made 25 and 45 days after transplanting using 30 kg/ha rates in the dry season and 20 kg/ha rates in the wet season. In the dry-season experiment 92 entries yielded over 8,000 kg/ha, with 24 of them, including 19 IR8 line selections, producing over 9,000 kg/ha. IR8 occurred four times in the trial and averaged 9,071 kg/ha. The highest yielding entry was an IR8line selection which gave 9,812 kg/ha, and one plot of this entry yielded over lO,O()() kg/ha. Five other IRRI dwarf selections yielded over 9,000 kg/ha. IRS and Taichung (Native) I yielded 8,503 kg/ha and 8,663 kg/ha, respectively (Tables 3 and 4). All the entries which yielded over 8,000 kg/ha were semidwarf improved plant-type selections. Yields in the wet-season experiment were reduced by wind damage including lodging caused by heavy rains during the ripening period. The highest yielding entry was IR8 which occurred six times in the trial and averaged 5,900 kg/ha. Eleven other sernidwarf selections from different crosses yielded over 5,500 kg/ha. Forty-nine selections yielded over 5,000 kg/ha. The promising selections entered in the two yield trials are listed in Table 3. This table also shows the number of selections and the yields (lowest, highest, and mean) of the selections of each cross, The yields of IR8, IRS, Taichung (Na tive) I and three popular Philippine varie ties are also given in the table for comparison. The highest total production from two crops

66

Varietal Improvement

D,ifY W,~t

Yl,a&Ciii1seasoll'll

H ~ghes:t Lowest

Mean 4576, 4773:

5216;

~R5913 IIRS: x [(CP·SlO~ x

Na:h:ftQ Moo S~4~

~AnSa III 58'91 R

~I 666 R

Na:tiilll'l:g Moo S.4x ~IR8 It (IFI· Hlf5, x; Dellg!'!Q.wOOogl'!n ~

n[IIJ~~I
.:;1'

644,5 a50~ 728,91 78910

::rim
4190 48B1
49'71[1

153'55
!5!5,~O

Ifi8

!(,

'~f:et:aj5x Be:l'le 4872

5,26:8

Patmd ~ HCp·S'lOI}f2 x Si,gadls]i x [IP-eta/3 xlli U~JI1 ] ~1'I6~3 lAB 1< I~O~wnx >EllB·25) ilFl586 IAe:.x U~PI':7i6J2 x T. ~172'~' ~'R578i IRS 1C" [Sigadiis,,xl"~~ll ~.~ :2 11'11127 'I'CP'-8LC)) x Si'9~is
mStl1 1R305 I A~8: )Ii (eN;; LO )! S~gad~;5ll 11 Si~i~/2!l n~1111 11:2 IR,Sx [Yu~,aral x .T(N ~1] I[P'lnal,3:.:: 'f(lrNJ) tl ,)I IB5S9A~· 18;·11/2 ): T(INI ~11l 7 IIRS x [B'5B9A4.1&1/2 x i(N~n 8 G,am P<li/2 ",i(N~ 1 4 l[p,e'tat.3 x T(N)11] It TKM·6 ~, Pna x i1F(lM,j~ Petw2 >l" T(I~ '111

5n 9
1511001
4941 3696:
,4:6791

5764

8706

757:2'

B~:%J 8008 8823

7(lS':;! 7636

9!H2

8~33
588:2

52'33 4404 637'01 51560 ~562 59'1 1

537
11

4445 "U1i.tl!9 2998 413:87

2654
41 1 1

3967

41 011 ,4837 4ge9'

lASS'
U'l600

4422
:53;23

489'7 4809!
470i7 3734 4361 54110 3767
4470

SJIH

61295 573:3
4761

sn.lo
8951 19 1
2'

1608.2'
81~3,

1:6:81
:2'706

~IR2!5:3
~IFl53:2 ~IR391
~I l9!5, f

8647
80913

9097 -S887 842~

8517
7925,

00\30
~Q6 80'14

51'[19

IIIR:2o:tJ:

Pl!ta/3 x n~,1 ~I
P!lHal4 x nMI1

3:
2

76013
;9031

5:270
4S:7i5

IIIR400
IIHS

8304 9071
8005

2:373 36!69 4059

4467 5915,

IIIAS

,44918
2i60~ ,4(122~ 272'2 3774

Peta
"'~N~'1 BP'I~:l!IG"l

C~·63
waS ,obl3~ k:g~. Wt. nUll, :ain~ lit <m:Jot!l1iI~d til' 15,9 W"1

61.5!03 86'63 6724 7478

PIO'mt ,htl:U;lklgy ~p~:r lff:l;elll~and f~p~eSl!(llI~ the ~f!C:eini~ of L'UJllpo·mf~cll;e!~5ecid;lw~~ a:rt~r rrJeiqg WI!!iddlti:~ I~Q y.i.eJd., dam Magrollomich'@!:tS. i foed IiIPQ~·byv~1IflB'm1!l$ iF!rcc~ lealiboppen. The wlieiWCrewl<Uilce.,omdmiU:ing and ,eootiJ'lg ,~U!a1ity reiliWngs: fo~liIa.cleriaJIlle~:f blight re&is,m:.n;Qe weJ{: ofth~ gl'al:ns wel'~ob'tSitn¢d10!l a.l!llhe~lltncsiJl,~ 6tl't!liilEled ~y 'thefts-s, l!I!a:f p1iI.f!iot"l.l~m(ltho~ land C[~ded m Ire yiehl trials, Detailed m~om1at~onl)!!l ~ose fOlf' bllc~er.i3illleaf shieak We~e bilsee!oll r.eW!ll! J8 Ciilu[~S rus g1vi~1'I irn, 'Fable: 4. DA!; d:a!ta O:ntIWl1ig;l'O obs~rv~t~Qtl~ ~tw~B~~,Th~ d;i!ltaofibl,af;t rc..:s:'iM .. [feIlCti,OJ]: '\\Va!S obtria;Jed from. greenhouse te:s:ts~y aJ]Icereplruseol ~ seed~!i:ng :rea.ctlo:H: to bl,as[ inlhe 67

"Ll!

OOOO<OOON

"':aiO"':aiai
L!lML!l~

....

<00

ait'i
m

000

aid

000"':0 .... _._ o:;io:;iai

L!lMN

-_

<tL!l..,.M<t

cciOOOO0:5

00

r.:

00

....

en
o ai

<0

<Xi
0«1

oo-oo,_v

~~~?ti;;;;:;
_j_j

"":00
NN

00

o:;i
N

~~~~g
_J: J:
_ _j_j_j _

~..,.<t""'N

omo
NMM

0000

_j

_j

_j

L!lmN<O<ON

~$;::~$~

00<0

........
00 ....

<'i..t N--=

«>N

",:..t

........

<'i

<t

....

00

o ....
o
M
<0

..... <t<tWN -.ir-iLri.._r",,: i""--r--..,....,. ...........

~~~*~;::
~<OM~OOO

<om..,.

L!l<O 00 <00 «>0 0>0 «> 0>

OOo<riaiai

.........

IOMN

""L!lw<t<t

<'i<rio

~ In

In

<Xl

00

'"' '"

..
.
UlUlUlUlUlUl

UlCI: CI:

a: a:: a:: a:: a::

Ul

a:: a:: a::

,",-

00-

..

UlUl UlUl

Ul Ul UlUl

Ul Ul UlUl

Ul

UlI

Ul Ul

<.J,D

Ul

Ul

(j}a::

Ul

o C,
C :::> <0

f-

0> 0<0 <0 000000

00

<ON

OJ<t

00'::

<t ....

00

'"
M

....

10

00

00 00 00

o o

MO

....
o

o o o

00000

.....

1000 OJW

L!l<O

010

OJ

00

'"

0000

<0 .... NO>OOO>

..... _

- ---m~
o
N

N"' ..... "'.....

00
N

O~N~MN _1'-" __

m~

m-

<0

m .... M<O

m_

O~

.... 0>«>

MNMMM 000>00

"'<t<O O~O <0

NO«>«>OO O«>~NN~ _~ __

-':-111"""

«>M «10

N m

0«> mo

«>«> 0«1

<0
«I

«> .... mo>

m
M

00

'"

m
mM
NNNN_

~~~~~g
OO~~~N M<tMM<tM

--;----

01

-- ----

..... IOO>

-.----~-

MN

woo
O>M
-OJ

0
N N

M<t

~m

MO
"'M

~
N

~N

~~
"'0

L!l

<t
M

O>N L!lM

o r--. M

01 MO M 00L!l .... L!l MOl o ML!lL!l NM <t O<t

~m

00L!l

.... <0 ..,.L!l .... 0

<ON

<0

L!l<t ....0 0

0>
N OJ

....J O L!l..... 0000

.~ .,
Z

0>

68

Varietal lmprovement

b1ast

IlIlJr'SeIi)'.

Brown fiw grains f[!om both !crop

ieiaJS'OH we~!f! at.laly!zed $or :pro~,e:incQ1illl!n~; y the S b

Cer~aI, (:h:eIil1:iSlry De~!!~ftrn.~n~"

IInt,ernatiiOna'11 'CoolperartivlJ irmi 1119 PlrogrSlitl

1

Variety

Uta and UtS b1. the d r:y' season, SCYCfSiI,p!romisiflg se:'lectiolls W'e.e P~aJ"l~d io~ seed :n:n:dli:p:l:ica1iollll. Eighteen of '1lI1~se seleetions came rrom [Pe:taj) x Jaiic)ung (NatiVie) 1]1 .x. l'KM-6, crOCSlS (lRSJ.l). fOllf from NahngMon S4 x Ta.lc:lillmg (N;atv.e) ~ (IRm60) aad
O.!l,c 1!';l'lclil: l:ro'lII1 Gamhj.12 lit TaialilU1nrg (NratiVoe) 1 OR253)j Cp·SW x MilS, (~R140).,;'[-76 X

Seed f\U:fiificali:on ,ilnd M!I!ltiiplli~ltiolll bi addi'Ltofl ttl prod!ucmg 'bNde:r seed. of

All: impo!liibnt pa:rt of tiJle ifl~!1lia~Oifiidp.rQgm.nl of 'Ille blStJiU~llieis d!.e supply:ing Qf seeds of breeding fules and 'lIad!!ti~5 .reqlil!estcd. by Icoope~a!ton .md ;iomru:S'ted o~,Elaniz;L'ltioln;, ,<l.neJ ;ind:iv~d~lals in '!he YlIfI']}@'I!S, rice·growing counttie~, DUrmg the yeilif, (jve. 1.2.000 samp:lcs of seeds
,Ormlls:titl.lte In',eedi~gline:s were tentta, 1,83 org:;miza'Uons !ii:'id il:ldivtduals in '79 ,couliI'h'ie-s. A I.IUge IiIW111be;r'Qf brc-.edi:lIg Hiles we re sent ilo '00-

(CP~SLO' X .Pel!l!) OR:29'8). Pe~.a x Taichuflg (Na,live) ,1 (U139')" Peta/2 X Tai~l:1iu!'lg (Native) I. ,(lR9's, and Peta/4 x Taicehul~g 9Na.l:i.ve)1 I, (lR400). Th~ area pLrmwiID each selec:tioll vlllried ro :froffi o;ruNijx.t.em,~h '[0 one:.:fOurth of' iill hechl!l'~, TIl,!~ prual'lti:li1igs: praY ided~fI lopporlunity te sec tilite performance (If tne;s;e' seleetioms under fi~ld eeadiUOIl:IIl!.MrO«l'OWI:'. coou!;h seed was ,3V,3!ila:bl~ die: lI.t end of the season foil' le-slilillg dI_.e :sel'!l>C~.ed UIiI,es 'of dliis g;roop O(n. a III'rge ~le. Se'{cn ~e IRS32lincs" namely, U~S32~1-2:18. I1UJ::n~:2:n., WRS32E239. UU32E251, IR.S32;5420 lR.532E42.3r, ,DId IR:B2ES76. 'Wi:l1>i! cons1tLer~d: p'ro:m;isiog and 'We[e in~luied ~n the d mui.tip~.i.c-,~t~ol:'lprogram. i,l:Il tho wet seasca. Six add:i1;i~!i}nal lines of {bis CIlO~ - lRS32·1-33, IR5'J2~f;160" n~,532E2 W, lR53 2:E2H ,.IRS32E230 ~d lRSl2E:521- we;1ie i.nCiI.lIille'diin 'this p.rogrnlll.

i;lpcwatof,s in ,B~~ma", Cey~o[),. Col.o;mbia. ~iladja. lndl(lI1e..~EI. KQr~i).. MaI!I,ysia. and Paikislan. tRRl rice breeders mmntain dose ICOOl?eE3ti~, with :r.ice' Ineeders in eaeh (if these countries duough a :free e~cl:la:Tlgo of hl!eas and b",",dili'ig ]1il:lue~Dals,. Rice brecdll~ii frum m.i'i.nycolllnl'r!.e:s willa come U! 'the mtlstitu te .~ research rel]orw~ lind ~ho)~s S!!lect liin.es from Ill!iUhUe breed:i!flg iiil1:useries lind yietdl tfia'i plou aad take the m;a.tc'r:iws home 'w:ith diem
1.lIt the eeneleslon of tbeir trmfl[i1IS pmgrams .. :F«l'qu,cI1Uy. UJ.e:y 'mii:k;e, c;rO$6S 'b;{!t,'i\II€!'ie[lUillevarieties, flOOl t]ie:i:l' home COllI ntrles a:nd ~~s:tUu t~ brei{l;d)· inS tililes d~tiiQIl their t!rllill:ill1g PiOPili'iiim and take lll,e: FI and :P2 !iced$; wH:h them .. A larg,e coil~th:m of v:a:rielies and ~i:!!IieS WI!I:,e ~iJjt tova:rffio~s,coolile~iUi~ calJ]! tries to Idetem:lme their IIdlllptaibUlty ~ rosiiStIlIllC1!' t.oV;lliriO,U$ dj$e;li~~. Jo,w temperatufcs and/or pe.!i~ontli'lnceuliHi~reon-

or

didmu

At ~e!i!$t ,OIIH'!.fourth of a heetllf'(! was pilllr:tt'e'di '~o hund~edll.lIll,lis I:lIlc.ludil:1g these [tom (1M!' IR.S/:::! X 'e',ll,cb0:£ ~.es:e ~.3 Sel(llOtIDoIlS,., 3IIlIdpr~!!ly·;jOW ;p'lots, .Zcll:idn, ,C[lOS'S 'Were ~l1it ~,o Ea5t Pakistan, Ceylon. Oif each of tlhe:sc li!n,es we re gfOwn tD ~tll.TlIIle seed ~ta.lays:iia. ,M,(i :liIrI.dciIl,esia, ~:o de:t'erm:ine 1I:liI.e:i:r re-

of d!et:p w,atef. Dllring

tile ye'.u. $cv~[t,al

purmcali,on p'rogram. Head JlOW blocks of tbree (iC~:iCfI1 to bacteria] le'M bl.i~t. Ow,r 800 Ilm,l!!,were' waxy lines from Gam P,ai/2 x Tai,dill1ug l(Nati.\'cl I. :S~f1t to Tbai]ail1d Ito ks:t. their resistiJ:Jlcte' b')ihc ftiID:1ciy, IR2S,J.]6.~2.. IR.lS3·~6.~.J.2!. and .• tYngro Y11:'1I unde~ field ,~(l!lJ(l:Itiolilsin that S :lR253,4.1.2;.]. we're gJOWIii mlOII,.S, wiilh a buak &I!if!\d, C01J 11I~:ry. illlu::rclIse. One-te!luh to one·twelfth of a heetase was, Af.ilJlnery of ~54, 1'n:ilti.t1!ue jc~dmg lines, from b plalllte,d wi~h bulk seed only of the following ] 3 di ffeltmt ~(is~ifl wni,cnoliie {i.fllli.epllrents seJecdoIi!$: was bLasl~re~~t.Jnt was .pre·pMCd. Tile Imes 'jillditldedin this 'mlU!re'ry had b~:111 founa, to be: U't. 1,60-2:5:·]·1 " NIlfu.gI!; MOil 54 II 1i"'MClluUJ,g>(1~aili\'e) ~ J"e-sist:an~underL()s Baflos Icond:itiO'JI$. "w.~lve rcWRHS0.4;}'1.2,. N~IIL!iIgM:on g.,4l, II 1,aicJil!llllg O>lative), ] S!S tan t p<l1re:n.tvaJrie'lies and lil lne breed'ifl§ Jines U'!:41l0-s:.9·3-J.;, Nitlllm,g, MDn S4f2x 'T;JIiduiI:n:g ,('N~,IiI\ilc)' 1
~IU4Q.U6, (CP2J,1 x SIlIQ'l7) X.MM UU 27-SQ-J..ti·l. (CP2.;'U !( SId().17 :;;,Sij;;idi$ IIH27·So..t.:lO'. (CPlll!! SliO-I1)r:1; S~ai!!l 1.R.:286·:1.8~2·1,Petlli: IDiiw,n X. Ta:ichllJJ:lg (N,~~i,yC)U [R.5g6·UO-z..~ • .lR8 x 1~l!In9:A4-Ul,fI2 .)1 1,ai,c]mng

f])om Tha.~IIl:lld; we.re also ilildudedif.'iLIil.ell,ucscry. ~~dwnpl:eli' of l!helj;_nes and varle ttes (!TOm lhis :mJrsery wer,e :scnt to Ceyl,oll.. 'Colom.b.ia., GUyn:liiil. bad ia, Il:1done~ia. East, PaJtjs~m. !IlI'Id Tha.iiI;l:lld"

~i IRs·lU·l-3·3.htll ~ Tmpa: 'Rotan

~B.fi .. e)

These liilL~S aad li'B,de:t ies ~a,ve bti~n phlLfil. ~[I me.sc' led ~mJint'li'ies ill'l.tW blast til:iliditn,sl' wilU be ti!!ken. Si:ll.~'

these countries presumably have different races of blast fungus, the reaction of the lines and varieties to races different from those prevalent at Los Banos will be determined. It is hoped that improved plant-type lines with a broad spectrum of resistance to blast will be detected. Breeding for High Protein Content The breeding program aimed at evolving highyielding varieties with high protein content is being carried out under a grant to the Institute from the National Institute of Health, U.S.A. The Fl and F 3 generations of the hybrid populations of crosses between IR8 and six high-protein varieties were grown in 1968. Brown and milled rice grains from 1.290 individual F 3 plants were saved for ana1ysis by the Cereal Chemistry Department. These plants were selected from 253 F3 lines, most of which showed high protein content in the previous generation. As shown in Table 5, the F3 plants grown from high-protein Fl plants (11 percen t or above) averaged 11.9 percent while F 3 plants produced from low-protein Fl plants (10 percent or below) averaged 8.8 percent in protein content. These figures represent only a part of the

1,290 plants saved as the rest of the data is still being analyzed. As indicated in Table 6, much of the increase in protein content is retained in the milled rice. The yield potential of these lines is not known. Most of the plants saved were from rows carrying the semi dwarf trait of IR8. At least some of the lines showing higher than average protein content were rated as having reasonably good plant-type. The protein content of most of tho entries grown in replicated yield trials and in observational yield trials was also determined. A comparison of data from the wet and dry season crops showed that protein content was lower during the dry season. The average yields and protein content of the hybrid lines of four crosses are summarized in Table 7. Most of these lines were high-yielding and averaged over 9 percent protein. Sixty-six individual good plant-type Peta x Taichung (Native) 1 backcross lines with a mean yield of 5.001 kg/ha averaged 9.1 percent in protein con tent (Table 8). The average protein content of 30 individual plots yielding over 5,500 kg/ha was 9.9 percent and of 7 plots yielding over 6,000 kgfha, 10.2 percent. Rough rice samples from a variety-fertilizer experiment conducted at The Ma1igaya Rice

Table 5. Protein content of F3 and F4 briwn rice grains from IR8 x high-protein varieties.'" Protein content (wet basis) 11% or over in F 3 brown rice gra ins F3 grains F4 grains Cross No. of plants Ave. protein content 12.1 11.7 11.2 11.6 11.3 13.1 11.8 53 237 No. of plants (%) 30 20 6 50 43 Ave. protein content 12.1 11.6 12.4 11.9 1·1.7 11.9 11.9 38 10% or below in brown rice grains F3 grains F4 grains No. of plants (%) 7 9 7 Ave. No. of protein plants content (%) 7.8 7.5 7.1 7.1 8.1 7.4 7.5 169 40 25 39 29 3 33 Ave. protein content (%) 8.6 8.7 8.7 9.0 9.0 9.2 8.8

IR8 IRB IR8 I R8

x x x x

Rikuto Norin 20 9 Omirt 39 9 Santo Chow Sung 10 12 12

4
5 6

IR8 x Crythroceros Korn. I R8 x Chok-jve-bi-chal Average Total

88

* The protein content of the parent variety check rows were: lAB, 7.8 (dry season] and 8.7 (wet season); Rikuto Norin 20, 9.5 and 12.9; Omirt 39, 9.B and 11.4; Santo, 8.8 and 13.1; Chow Sung, 9.8 and 10.8; Korn., 9.7 and 12.2; and Chok-jye-bi-chal, 10.0 and 14.5. Crythroceros

70

Varietal Improvement

Table 6. Comparative protein content of brown and milled rice of F 3 and F q caryopsis from crossed plants of IR8 x high-protein varieties.

Table 8. Average yield and protein content (wet basis) of hybrid lines IRRI, I %8 wet season (observational yield trial),

Protein content (% wet basis)

Average IR No. IR265 Cross Ch. 242/2 x (PI 215936 x Mo. R·500) I R95 IR262 IR400 I R482 IR564 IA658 Peta/2 x T (N) 1 Peta/3xT(N)1 Peta/4xT(N)1 Peta/5 x T(N) 1 Peta/6 x T(N)1 Peta/7 x TIN) 1 8 10 14 7 9 18 5308 5555 5266 4461 5450 5435 9.3 8.9 9.3 8.9 9.3 8.9 No. of lines 12 Yield (kg/ha) 4509 Protein content 8.2

F3 caryopsis
Brown rice

F4 caryopsis -B-ro-w-n--M-il-Ie-d

rice rice ------------------------------10.9 High-protein plants 11.8 11.9 ( 11% or over) Low-protein plants (10%orbelow) IR8 7.5 8.8 8.7 8.3 8.1

Table 7. Average yield and protein content (wet basis) of hybrid lines, IRRI, 11)68 wet season (observational yield trial), *

IR No.

Cross

Average No. of Yield lines (kg/ha) 19 18 11 13 4284 5325 4598 4977

Breeding Program with TKM-6

Protein content 10.3 9.1 10.9 8.8

IR532 IR580 IR751 IR661

[Peta/3 x T(N)1] x TKM·6 IR8xTKM-6 I R8 x (Peta/5 x Belle Patna) I R8 x {CP-SLQ x Sigadis

* 30 plots averaging over 5,500 kg/ha averaged 9.9% protein (range 8.5·11.3%). 7 plots averaging over 6,000 kg/ha averaged 10.2% protein (range 9.4-11.0%).

Experiment and Training Center in Nueva Ecija, Philippines, by the Agronomy Department was milled by the Varietal Improvement Department and analyzed for protein content by the Cereal Chemistry Department. The protein content of both the brown and milled rice of four improved varieties (IRS, IRS, IRS32E576 and C4-63) and Peta increased substantially as the fertilizer rates increased (Table 9). Likewise, head and total milled rice yields tended to increase with higher levels of nitrogen.

The variety TKM-6 from India was found to be highly resistant to stem borers by Institute entomologists, and resistant to bacterial leaf blight and the tungro virus by Institute plant pathologists. Because of its insect and disease resistance and good grain quality, TKM-6 was considered a potentially good parent for the breeding program. In late 1965, plant breeders crossed TKM-6 with a dwarf selection from Peta/3 x Taichung (Native) I. An F2 population was grown in March 1966 and selected plants were saved to form an F 3 bulk population which was planted in November 1966. A total of 255 promising plants were selected from this F3 bulk population and planted to F4 pedigree rows in May 1967. The F sand F (, pedigree lines were grown in pedigree nurseries planted in Novem ber 1967 and May 1968. The selection for plant type, disease reaction, grain quality, and maturity date was continued. A part of the hybrid material emanating from the above cross was also evaluated cooperatively by the Entomology Department. The entomologists made many panicle selections from the F3 bulk populations grown by the Varietal Improvement Department and grew several F4 bulk populations in March 1967. The over 600 Fs plant selections made from these populations were planted in August 1967 in two-row plots which were replicated twice. One replication - was pro-

Vane tal Improvement

71

Table 9. Protein content (wet basis) of brown and milled rice, rough rice and milling yields from a fertilizer experiment, MaJigays Center, 1968 wet season.

Nitrogen late {kg/hal Variety I mproved varieties ~ Protein content Brown Milled Rough rice, kg/ha Head rice. kg/ha Total milled rice, kg/ha Peta Protein content Brown Mi.lled Rough rice, kg/ha Head rice, kg/ha Total milled rice. kg/ha .. Average of four varieties

30

60

90

120

150

7.09 6.22 3854 2090 2666

7.52 6.58 4406 2469 3061

8.01 7.25 5180 2989 3634

8.36 7.43 5206 3083 3629

8.47 7.74 5710 3438 4042

9.48 8.47 5337 3261 3777

7.07 6.20 4110 2085 2879

7.40 6.70 4334 2306 3022

8.10 6.65 4280 2455 3019

8.70 7.50 4572 2773 3200

8.87 7.80 4539 2728 3256

8.03 7.63 4745 2918 3386

{I 8,IR 5,C4·63 and IR532E576). R

trials grown in December 1967 an d June 1968. Twenty-seven lines, 17 from the Entomology Department nursery and 10 from the Varietal Improvement Department nursery, were included in the dry-season yield trial. At least four lines yielded as well as IR8. The wet-season yield trial planted in June 1968 included 19 lines from this cross. The average yield of these lines was much lower than those of IR8 and other high-yielding entries (Table 3) mainly because most of the lines lodged badly. During the 1968 dry season, lines 0 f this cross which were included in the seed multiplication program were evaluated on a field scale. The seeds of three of the more promising lines, namely IR532E239, lR532E257 and IRS32E576, were distributed to cooperators of the Philippine Agricultural Productivity Commission for testing in 13 provinces of the Philippines during the wet season. The Agronomy Department carried out fertility trials with these lines at four locations in the Philippines during the 1968 wet season. The Varietal Improvement Department again multiplied the seeds in the 1968 wet season and at the end of that season enough seeds were available for

tected with insecticides while the other was not. In the unprotected plots, entomologists classified the selections as to the degree of stem borer resistance, while in the protected plots, plant breeders classified the lines as to plant type, disease reaction, and other agronomic traits. Sixteen promising lines good plant type and of a low level of stem borer infestation were harvested and then grown as a part of the seed multiplication program. Single plant selections from the above pedigree nursery were grown in January 1968 in F6 pedigree rows in two replication - one protected, and the other unprotected. Again, the material was studied jointly by the entomologists and plant breeders and selections from this nursery numbering 621 were grown in the pedigree nursery of the Varietal Improvement Department in June 1968. All lines were then evaluated for grain dormancy, threshability, and resistance to bacterial leaf blight, bacterial leaf streak, blast, and the tungro virus. Data on grain tex ture, size and shape, milling recovery, amylose content and gelatimization temperature were also obtained. The yielding ability of the lines of this cross was tested in the observational and replicated yield

Varietal Improvement

sending to other countries for testing under local conditions. Seeds of IR532E576 which appeared to be one of the best lines were sent to Burma, Colombia, Ceylon, India, Indonesia, and Pakistan for large-scale testing. Small samples of this selection have been sent to many other countries. At the end of the 1969 dry season, data on the international performance of IR532E576 will become available. Meanwhile, the purification of seed of several lines which was started in the 1968 wet season is being continued; if any of the lines are recommended for cultivation, breeder's seed wiJI be made available. The promising lines of this cross have the following main features: They are moderately resistant to bacterial and virus diseases, and some lines are resistant to blast under Los Banos conditions. Their level of resistance to stem borers is not as high as that of TKM-6 but is considerably better than that of the other commonly cultivated varieties. They have fine grains of clear texture with high amylose content and are dry cooking. Their maturity period varies from 20 to 25 days earlier to a few days later than that of IR8. Most of them have good plant types with good early vegetative vigor. However, all of them have weak stems and may lodge with heavy nitrogen fertilization. IR580, a subsequently lines of this but the grain cross between IR8 and TKM-6, was handled in the same manner. The cross have somewhat sturdier stems quality is generally poor.

Strain

Culm length (em) Origin 28 35 47 43 45 Japan China France Taiwan IRRI U.S. U.S. Taiwan IRRI U.S.

Parentage

Daikoku dwarf (dl) Ai-yeh-lu dwarf Fanny sernidwarf FF36 IR273

Intermediate dwarf 69 Long-grain dwarf 66 Taichung (Native) 1 58 IR8 Ace. 6993 63 67

[(Chianung 242.x Taichung (Native) 1) x Taman 31 (CP231 x SLO-17) /2 x Taichung (Native) I F4 Dee-gwo-woo-gen x Tsai-yuan-chung
Pe ta x Dee-ge o-

woo-gen CP231 x SLO-17

Genetic and Cytogenetic Studies

Allelic relationships of dwarfs and semidwarfs In the Institute's breeding program, the recessive gene in Taiwan's sernidwarf varieties [Dee-geowoo-gen, I-geo-tze and Taichung (Native) 1] has served as the principal source of short plant stature. Another promising source of sernidwarfism came from the CP231 x SW-17 selections (Ace. 6993 and sister lines). Samples of several other sources of sernidwarfisrn and dwarfism were taken from the Institute's world collection in an attempt to broaden the genetic base of potential breeding material. The following 10 short-statuted lines were inclu ded in this genetic study.

The 10 short-statu red selections were intercrossed. Each line was also crossed with two tall tropical varieties, BPI-76-1 and Peta, Among the F 1 hybrids and parents grown in the 1967 wet season, most of the F 1 plan ts originating from dwarf x dwarf, dwarf x semidwarf, and semidwarf x semidwarf crosses exceeded in height the tall parent involved in the crosses. But only a few of these hybrids were almost as tall as the tall tropical Varieties. On the other hand, the crosses between '3 tall parent and a short-statured line produced Fl hybrids which were shorter than the tall parent, indicating the incomplete dominance of tallness. Sixty-four F2 populations and their parents were grown in the 1968 dry season. About 200 plants were included in each F2 sample. The cool temperatures which prevailed in the first part of the crop season adversely affected the full expression of the height genes. For instance, the average culm length of Peta plants reached only 114 ern, while the culm length of BPI-76-1 was only 83 em, From crosses among the dwarfs and semidwarfs, the F 1 distri bu tions indicate tha t the following groups of strains can be recognized as having non-allelic gene or genes for short stature. I. Daikoku dwarf and Ai-yeh-lu dwarf. 2. Fanny semidwarf. 3. Intermediate semidwarf and .Jong-grain semidwarf. 4. Accession 6993 and IR273. 5. FF36, Taichung (Native) 1, IR8 and IR273.

Varietal Improvement

73

The F2 data from dwarfx tall and semidwarfx tall crosses indica te that: I. The dwarf stature in Daikoku and Ai-yeh-Iu lines is controUed mainly by a single recessive gene and probably involves genes of the modifying type (modifiers). 2. Sernidwarfism in the Fanny strain is conditioned by genes of a polygenic-additive nature. 3. The intermediate and long-grain semidwarfs carry a number of recessive alleles for short stature. 4. FF36 has the recessive gene from Taichung (Native) 1 (originally from Dee-geo-woo-gen) and other modifiers for short height. This recessive gene is also present in IR8. 5. The IR273 line combines the polygenicadditive genes from Acc. 6993 and the recessive gene from Taichung (Native) 1. In addition to the above generalized postulates, most of the F2 populations from dwarf x dwarf and dwarf x semid warf crosses involving non-allelic genes did not produce segregates which were as tall as Peta. On the other hand, the dwarf x semidwarf crosses frequently yielded extremely short F 2 progenies. Similarly, the semidwarf x sernidwarf crosses involving non-allelic genes produced very few tall segregates. Crosses involving the intermediate and long-grain sernidwarfs frequently produced a few F2 plants which belong to the dwarf class. In contrast to the above, crosses between Peta and the Ai-yeh-lu dwarf or either one of the U.S. semidwarfs (intermediate or long-grain) produced segregates which were taller than Peta. The above three short-statuted selections tended to produce relatively taller segregates in their crosses with other sernidwarfs, The diverse F 2 distribu tions found in the 64 crosses suggest that: 1. Tall plant height is generally dominant over short stature, but the dominance is incomplete in nearly all cases. 2. A complex of height genes is involved in the crosses, These genes vary in their degree of domino ance, magni tude of effect, and level of heritability. The modifiers also differ in the direction of effects (enhancing or depressing). 3. The Fanny semidwarf, FF36, Ace. 6993 and Taichung (Native) I, appear to have in common several negative modifiers (for short stature) which

could explain the failure to recover tall segregates in their F 2 samples, 4. The expression of height genes could be markedly affected by the different genes for maturity which were also segregating in these crosses, especially those genes for extreme earliness carried by the dwarfs and some of the semidwarfs.

Inheritance of photoperiod sensitivity Observations made on F2 samples of three photoperiod-sensitive x sensitive crosses were given in the 1967 Annual Report. The remaining three crosses from this series were planted in late May under natural daylength and they involved Podiwi-A (8) of Ceylon as the common parent. The other parents were BPI-76, Siam 29 and Raminad Strain 3. The four parents were also included in the field planting. The mean duration from seeding to heading ranged among the parents from J 33 days for BPI-76, 166 days for Siam 29, 184 days for Rarninad Strain 3, and 188 days for Podiwi-A (8). The parental populations of about 100 plants each headed within a period of 6 to 8 days. The three F 2 populations showed different distribu tion curves bu t indicated three common features. 1. AI! of the F2 plants could be classified as photoperiod-sensitive under a natural daylength. 2. The total range of F2 distribution was related :0 the magnitude of difference between the parents concerned. 3. Only a few F2 plants flowered later than the Podiwi parent, whereas the earliest F2 plants headed at the same time as BPI-76. The above observations are similar to those made on the 1967 wet-season crop. The field da ta again suggest that the major photoperiodsensitivity gene (Se) in the four parents may belong to a multiple allelic series or a complex locus. In order to have a fuller description of the genes concerned, it appears necessary to use controlled photoperiods and vegetative tillers of the same plant so that concurrent determination of genes controlling the components of photoperiod sensitivity (the critical photoperiod and the optimum photoperiod) could be made.

74

Varietal Improvement

Cytological study of sterility in indica x indica and indica x bulu hvbrids"

Studies on the sterility of intervarietal crosses were continued in 1968. Twelve indica x indica, eight indica x bulu, and one bulu x bulu crosses involving 10 indica and 2 bulu varieties were used. The spikelet fertility of the parent varieties ranged from 51.5 percent in TKM-6 to 90.3 percent in IR8. Among the F 1 hybrids, fertility ranged from 21.1 percent in Boegi imba x Basmati 370 to 85.1 percent in Pankhari 203 x Basmati 370. All the parent varieties showed essentially normal chromosome behavior at meiosis except for a low frequency of univalents (0.9 to 2.8%) in three parent Varieties, 2 IR's (1.5%), laggards (1.5%), and a bridge (1.5%) in three other varieties. The hybrids exhibited a wide range of chromosomal aberrations. At diakinesis and metaphase I, the following abnormalities with their corresponding frequency were observed in some of the hybrids: 1 to 2 IV's (0.3-4.2%), 2 to 4 I's (0.2-4.3%), a ring-of-four chromosomes (0.3·0.7%), 2UI's (0.8%), 1 III + 1 I (0.2%·1.0%) and a persistent nucleolus at metaphase I (0.2%). At anaphase I and telophase I, laggards (0.4-2.6%), a bridge withou t fragment

(0.2-0.9%), and a bridge with fragment (1.0%) were also observed in some hybrids. There was no significant difference in kind and frequency of the abnormalities observed between the indica x indica and indica x bulu hybrids. Although some abnormalities observed in the hybrids were not found in the parents, their frequency was too low to accoun t for the observed sterility. Furthermore, there was no association between sterility and kind or frequency of abnormalities observed, e.g., the most sterile hybrids had only 2.8 and 0.6 percent abnormalities at diakinesis - metaphase I and anaphase 1telophase I, respectively. The most fertile hybrids had 2.3 and 0.6 percent abnormalities at the same stages. The data suggest that sterility in indica x indica and indica x bulu hybrids may not be due to chromosomal aberrations alone. These observations and conclusions are similar to those on the indica x japonica hybrids. As in the indica x japonica hybrid, sterility could be attributed to more complex factors such as genic imbalance in addition to chromosomal aberrations.

• A cooperative project between lRRl and the College of Agriculture of the University of the Philippines..

Varietal Improvement

7S

Plant Pathology ..
OJ

<:

.9-

[1

The broad spectrum of resistance to the blast disease of several rice varieties was confirmed by additional tests in the Philippines and in various countries participating in the International Blast Nursery program. Experiments were initiated to determine whether rice varieties possess horizontal (field) resistance to blast. Many more new pathogenic races of the fungus were characterized in the Philippines. and the presence of many and new races in the Institute blast nursery was demonstrated. These and other experiments substantiated the concept that great variability exists in Pyricularia oryzae. The varietal resistance to the bacterial blight disease of selected varieties was further investigated. Experiments showed that bacterial blight is not transmitted through seed. Many varieties were found resistant to the bacterial streak disease. Information was also gathered on the virulence of the bacterial strains, bacteriophages, host range, etc. of bacterial streak. Two new vectors which transmit the tungro virus were found. The morphological features of the virus transmitters and nontransmitters of both Nephotettix apicalis and Nephotettix impicticeps were studied in detail. The extensive host range of tungro virus was also determined, with the wild rices appearing to be perhaps the most important virus reservoir during the off-season. Strains or biotypes of the tungro virus vector which can persist and develop on resistant varieties were isolated. Resistance to the tungro virus seemed to be distinct at the seedling and mature plant stages of rice varieties. A mass screening method for grassy stunt disease was developed, and several thousand varieties and breeding lines were tested, with not one proving to be highly resistant. The transmission of grassy stunt was investigated in detail.

77

00

001)

001

00000

0000'1)01)

s~

9gg83g83
ClIO 000 0

2
I)

i
•

78

Plant Pathology

Rice Blast Disease

Varietal resistance The screening for resistance to the rice blast disease in the International Uniform Blast Nurseries (IBN) during the last several years has so far yielded 159 test results in 24 countries. This is the first time that rice varieties have been tested over such a wide geographic area and for many seasons, and it has made possible a more comprehensive assessment of varietal reaction to the blast disease. The test varieties showed a full range of reaction from very resistant to very susceptible. Some showed a broad spectrum of resistance, susceptible only in very few of the tests, while others showed only a few resistant reactions. Of the first set of 258 varieties tested in the IBN, 18 were found to be most resistant and 2 most susceptible (Fig. 1). These resistant varieties are generally considered to be the most useful available sources of resistance to blast. The data in Fig. 1, as well as results of numerous artificial inoculations made in Japan, the United States, and the Philippines showed the variety Tetep to be the most resistant. Many other varieties are resistant in specific countries or geographic regions of Asia, such as Japan- Korea, Sou theast Asia, and IndiaPakistan. *

highly resistant varieties may also have some type of field resistance. Horizontal (non-race-specific or field) resistance to blast When varietal resistance is due to major genes and when a pathogenic race exists which can heavily infect the variety, it is feared that such resistance will break down after the variety has been ex tensively cultivated. It also was shown recently that the blast fungus is highly variable and can produce new races with high frequency. Experiments were therefore initiated to seek the horizontal or nonrace-specific type of resistance, with which a certain degree of resistance is maintained regardless of the pathogenic races. However, as horizontal resistance is polygenic, intangible, and difficult to assess in most cases, separation of field resistance from major gene resistance usually requires repeated tests over a long period and in various geographic areas. An international cooperative study should prove useful. There are two types of varietal reaction to blast which are tangible and may be considered as field resistance: (I) The small-lesion type (type 3), which is common in the blast nursery. This type of lesion produces few conidia (estimated at 15 to 50 each day for a few days) as compared to many in large lesions (4,000 - 6,000 each day for more than 2 weeks, Fig. 2). (2) The few-lesion type, in which typical lesions (type 4) are produced, but their number on each seedling is small in contrast to the 50 to 100 or more lesions on susceptible varieties. The small-lesion type of reaction, however, is in most cases controller! by major genes, as many of the varieties produce large lesions and become susceptible in subsequent tests. Whether varieties possess such reaction bu tare non-race-specific, i.e.., field-resistant, can only be determined by repeated tests over wide geographic areas. The few-lesion type of reaction in many cases is due to the low population of the specific race(s) present in the nursery. Many varieties reacted this way in the early blast nursery tests but became very susceptible in the later tests as the population of the specific race(s) increased. To determine

A second set of 321 varieties (referred to as "BRWCV Set" in the 1967 Annual Report) selected from over 8,000 entries in the Institute blast nursery has so far been subjected to 49 tests in 10 countries. Some of those considered as highly resistant are also listed in Fig. 1. This set of varieties will be tested further in the IBN. Since even the most resistant varieties were susceptible in one or more readings, it was generally assumed that resistance in these varieties was due to major genes. The IBN data are not detailed enough to indicate whether these varieties also possess some type of horizontal (field) resistance. For instance, the varieties in the blast nurseries with partly dead leaves are usually considered susceptible. This susceptibility, however, may have resulted from numerous type 3 lesions due to the very heavy inoculum in the nurseries. The type 3 reaction rna y be considered as a type of horizon tal resistance in the field. Therefore, some of the

Ri.ce Commission

• The results are published in detail in the International Newsletter 13 (3): 22-30, 15 (3): 1'10, 17 (in press).

PLant Pathology

79

Fig. 2. Conidia released from a typical blast lesion collected in one night on agar media.

whether this type of reaction is due to a low population of pathogenic race(s) or to field resistance, the fungus on these few lesions may be isolated and cultured to increase the conidial population, and then artificially inoculated on the varieties. If the variety consistently produces only a few lesions in spite of repeated inoculations, it may possess field resistance. Small-lesion type of resistance. A group of 441 varieties showing the small-lesion type of reaction in the early tests in the blast nursery was again tested. After the first repea ted te st, only 301 varieties remained resistant (type 3 or better). 0 f this total, 265 (there were not enough seeds for the other 36) we re again tested and 218 remained resistant. These varieties 'will be tested further in the IRRI blast nursery, in other cooperative nurseries in the Philippines and, in the near future, in the International Blast Nursery. Unless extensively tested, there is no way of determining whether this reaction (Type 3) can be attribu ted to major genes or to field resistance. Small number of lesions by artificial inoculation. Many varieties showing typical but few

lesions were found in the blast n-ursery. The fungi in these leaf lesions were isolated, cultured, and artificially inocula ted with heavy inoculum to the varieties at the seedling stage. Variety Khao-tehhaeng 17 (susceptible to most races known in the Philippines) was included in each inoculation for comparison. Preliminary results (Fig. 3,A) seemed to show that some of these varieties have fewer lesions than Khao-teh-haeng 17, although the fungi were isolated from them. Two of these varieties, however, have as many lesions as Khao-teh-haeng 17. Apparently, this scarcity of lesions in the blast nursery was due to the low population of the races which attacked the two varieties. It is also interesting to note that the varieties with fewer lesions were also those which were found to be more resistant in the IBN tests. Whether these varieties also possess this few-lesion type of field resistance is yet to be confirmed. In another set of experiments, the varieties Tetep and Khao-teh-haeng 17 were included in the artificial inoculations with the fungus isolated from the other varieties. Again, in many cases, more lesions were found on Khao-teh-haeng 17

80

Plant Pathology

~ 40~~~---~~~-~/S~

~~---~~

the field where the inoculum occurs repeatedly, and whether this type of resistance in leaves also exists in panicles, require further investigation. The reasons for the varying number of lesions on Khao- teh -haeng 17 in di ffere n t in ocu lations are not definitely known. In some cases, the variability was due to the concentration of the conidial suspension used in the inoculation since some isolates produced fewer conidia in culture although, in most of the cases, this was adjusted to a similar concentration. Also, it m ight have been partly due to the fluctuating environmental conditions in the greenhouse which cannot adequately be controlled. Small number of lesions in blast nursery. Eight rice varieties from the blast nursery were selected to study the rate of blast disease development by counting the number of lesions observed every other day on the seedlings in 50-em rows from the time the disease first appeared. The susceptible check variety, Tjerernas, was olanted in parallel 10
~
KTH-17

o~
Or iginal Voriet ie s

Fig. 3,A. Numbers of blast lesions developed by artificial inoculation on leaves of variety Khao-teh-haeng 17 (KTH 17) and on those of original varieties, from which the fungus was isolated.

80

1--------+----

.......... r---1

than on the original varieties; in some cases, the original varieties showed as many lesions as Khaoteh-haeng 17. In all cases, however, Tetep had the fewest lesions (Fig. 3,8), and these were mostly of type 3. As mentioned earlier, Tetep is the most resistant variety found so far in the IBN. More significant differences were found by comparing the number of type 4 lesions found among the varieties. When varieties produce only a few lesions even with the fungus isolated from them, i.e., pathogenic to the varieties, and many lesions are produced on the susceptible control variety, Khao.teh-haeng 17, in the same inoculation, it is possible that these varieties possess some kind of field resistance. Previous studies at the Institute have shown that in a culture, even that from a single conidium, several races may be present. Some of the races may not infect the varieties from which the culture originated, hence, the lesions would be few. Whether these varieties possess some kind of field resistance, whether they will be resistant in

"E

0:
<,

<:>

60

"' '" .3
c: .2

'0
Z

s
E

40

20

Tetep

OrjQinol Varje1ies

KTH-17

Fig. 3,D. Numbers of blast lesions (type, 3 and 4) developed by artificial inoculation at the same times on leaves of varieties Khao-teh-haeng 17, on Tetep, and on the original varieties from which the fungus was isolated.

Plant Pathology

81

4000

'2000

•
0

Tjere Mas
H-5

1000

,
v

Tetep
Tadukan PI 180061

500

bably the result of the inability of many of the races present in the blast nursery to infect these varieties (major gene resistance). Since there are numerous fungus races and almost all the rice varieties have resistance to a few or many of them, it is difficult to separate horizontal resistance from resistance due to major genes. Nevertheless, these resistant varieties greatly reduce the effective inoculum, thus minimizing disease epidemics.

o ';;;

~ 200

Pathogenic races in the IRRI blast nursery

iii
'0
:::I

~ 100

Z
E
Z

50

20

10

3rd

5th Reading

7th

9th

Days

Fig. 3,e. Number of blast lesions observed on four more resistant varieties and their neighboring susceptible checks in blast nursery (100 seedlings).

em away from each of the varieties for comparison. The results showed that all the 8 selected resistant varieties had much fewer lesions than the neighboring check variety. The data from four of the eight varieties together with the check variety are plotted in Fig. 3,e to illustrate these differences in lesion number. The number of lesions (both type 3 and type 4) was counted on the basis of 100 seedlings. Superficially, it appears as if the eight varieties have considerable field resistance. However, the small number of lesions on the varieties was pro-

It is generally assumed that several races of the blast fungus may be present in rice fields. However, the number of these races, their relative population, seasonal changes, etc. have not been investigated in detail. A study of these aspects was conducted with samples collected from the Institute blast nursery. A total of 363 samples was collected monthly and monoconidial cultures were obtained during a period of 21 months. Each isolate was inoculated to 8 international and 12 Philippine differential varieties so that both the international race groups and the Philippine races to which they belong could be identified. The 363 isolates were differentiated into 37 international race groups, of which 22 were new. Of the 60 Philippine races, 34 were new. More races would have been detected had more samples been taken. The number of races, as indicated by the samples, varied from 3 to 14 each month with an average of about 9. The composition of races differed each month, and the race groups in any two months were not similar. The relative population of the races detected in each month also varied; a race may have a high frequency (percentage of total isolates) in one month but a low one in other months. A few races were present in most of the months, such as IA-109 (20 in 21 months) and P-8 (19 in 21 months), and showed high frequency; hence they were considered as the prevailing or major races. Other races occurred only periodically, and many were detected only once during the entire period. These changes are illustrated in Fig. 4. The relatively large number of races detected was probably due to the presence of a very susceptible variety and of several hundred other varieties in the blast nursery. We believe that most of the new races are being produced by the fungus

82

Plant Pathology

100

90 • 80 P-8

0
.~

'" "0

<f>

70

0Iill P-12
~ ~ P-19 P-20

P-9

-0

§ 60

'0 CO

II P-30

a. 50

P-36

'"

c .2

0 40

~
-", Il:
OJ

<i3

"0

30

20

10

A S 1966

J F 1967

J F 1968

Fig. 4. Races of Pyricularia oryzae Cav, detected in lRRl blast nursery each month from August 1966 to May 1968 and their relative population (based upon Philippine differentials).

in the blast nursery rather than by air-borne conidia from outside. A larger number of races than those detected may actually have been produced.
New pathogenic races in the Philippines

Up to 1967, 88 races have been characterized by the Philippine differential varieties and 49 races by the international differential varieties in the Philippines. In 1968, 28 new Philippine races and 18 new international races were identified. Their designated numbers and varietal reactions are shown in Tables 1 and 2.
Variation in pathogenicity from single cells

Our early studies have shown that many pathogenic races may be differentiated among conidia produced on single lesions, and may be produced by subcultures from a single conidium. Of 50 sub-

cultures tested, one culture (L-2-36), however, did not change in pathogenicity. The variability and stability of the fungus were further investigated. Variation in pathogenicity among subcultures of Single cells. Single hypha! tips of two germinating conidia from cui lure 1-42 (a variable culture) were isolated from both the apical and basal cells of each of the conidia. These were designated as A-a (Conidia A, apical cell), A-b (Conidia A, basal cell), B-a and B-b. They were cultured and 15 to 20 of them were further isolated from each of the subcultures. These were grown and the conidia collected and inoculated on the international and Philippine differential varieties. Based on the international differential varieties, the 20 single hyphal-tip isolates of A-a were separated into 3 races, and the 17 A-b isolates into 7 races, the 17 B-a isolates into 2 races, and the 15 B-b isolates into 5 races.

Plant Pathology

83

Table I. New races or Pyricularia oryzae in the Philippines

based upon international differentials PJ.180061 [Dular]

differentials.

International International race number IA-66 IA-67 IA-68 IA-97 IA-121 IA-124 IB-61 IC-26 IE-l IF-2 IF-3 P.1.231128 (Raminad 5tr.3) 5 5 5 5 5 5 R R R Zenith P.1.201902 (NP-125) Usen

Kanto 51 C.1.8970(s)

C.1.1561 (Caloro)

Number of isolates

R
R

R R R R R R 5 R R R R

R 5 5 R R R R 5 R

R
R

5 5 5 S R R R R R R R

5 S 5 S R R R R

5
R R

5 S S 5 S 5 R 5 S S 5

5 R R 5 5 R S 5 5 S R

R 5 R S S R S R

5
R S

2 9 1 3 1 1 1 1 1 1 1

Table 2. New physiologic

races of Pvricuteri« oryzae (identified Differential

in 1968) based upon Philippine differentials. varieties Peta

Philippine race number P·89 P·90 P-91 P-92 P-93 P-94 P-95 P·96 P-97 P-98 P-99 P-l00 P·l01 P-102 P-l03 P-104 P-l05 P-t06 P-l07 P·108 P-109 P-ll0 P-ll1 P-112 pot 13 pot t4 P·t15 P-l t6 P-l17 P-ll8

Kataktara Da-2

C.1.5309 Chokoto Wag-wag Co25 Paikantao

se. 3

Raminad

Taichung T.C.W.C.

Lacrosse Sha-tiao- Khao- No. tsao (s] teh. of haeng17 isolates

5 S 5 5 5 S R R R R R S R R R R R R R R R R S 5 5 5 5

5 5
S 5 S

5
R R R R

5
S R R S 5 S 5 S 5 S R 5

S 5 S 5 S 5 R R R R 5

5
S R S 5 R

5
5 S S

5 5
5

5 5
5

5
5 5 S

S 5 5 5 R 5 R R R R 5 R R S S 5 R R R R R R R R R

R R R R R R

5
R R R S R R R R R R R R R R R R R R R R R R R R R R R R R

5 5
S

5 5 R 5 S R R R R

5 5
R

s
R

5
S 5 R R 5 S R S 5 R R R R S

5
R R

5
R R R 5

5
R S R R R

5
S R R S R R R S 5

5
S R R R R R R R R 5

5
R R 5 R R 5 5

5 5 R 5 5 5 S R R R S R R S 5 5 5 S 5 S 5 5

5 S R S R 5 R R 5 5

S R R S R R

5
R R S

5
R 5 S

5
5 S

5
S

5
S

5
S 5 5 5

5
S

5
5 S S

S 5 S S 5 S 5 S S S S 5 S S S 5 5 S

5
S

5
S 5

5
5

5
S

5
5 5 5 5 R 5 5

5
S 5 5 5 S

5
S

5
R R R R

5 5
S 5 5 5

5
R S

5
5 5 S 5 S 5 5

5
R R

5
R

5
S

5
R

5
S

3 4 1 5 1 1 1 1 1 1 2 1 1 1 1 1 11 14 2 1 t 1 2 4 4 1 1 1 2

84

Plant Pathology

Based on the Philippine differential vaneties, the 20 A-a single hyphal-tip isolates were separated into 7 races, the 17 A-b isolates into 8 races, the 17 B-a isolates into 8 races, and the 15 B-b isolates into 8 races (Table 3). This further indicates that the fungus may produce new races not only from a single conidium, which is three-celled, but also from single hyphal tips isolated from single apical or basal cells on the conidium. Among the races, lD-9 or the original race, P-23, still dominated the population, i.e., it had a higher frequency. A total of 9 international race groups and 19 Philippine races was separated from the single hyphal tip isolates of the two conidia.

Variability and stability of monoconidial subcultures. Two isolates, 1-281 and L-2-36, were
studied further. Forty-one monoconidial subcultures from 1-281, when inoculated on the international and Philippine differential varieties, were separated into 3 international race groups and 4 Philippine races as follows: by international differentials: IA-65 (25 subcultures), IC-I 914), and ID-l (2); by Philippine differentials: P-8S (23), P-84 (IS), P-37 (2), and P-86 (I). Twenty-six monoconidial subcultures of L-2-36, which was known to be stable, were further tested on the two sets of differentials. All

of the subcultures had the same reaction and were identified as IA-I09 or P-8. The available results seem to show that certain isolates, such as L-2-36, are quite stable in pathogenicity. Some are variable but the frequency in which they produce new races is low, such as 1-281, while others, such as 1-43, are very variable. Variability in single lesions was further studied using Lesion No. S. When inoculated to the two sets of differentials the 24 monoconidial cultures isolated from the lesion were separated into 6 race groups: IA-6S, IA-67, IA-68, IA-124, lB-l and IC-3. Eleven of the 24 monoconidial cultures belonged to IA-6S, 9 to IA-6 7, and one each to the 4 other race groups. Based on the Philippine differentials, the 24 cultures were separated into 12 races: P-S8 (I culture), P·86 (I), P-89 (2), P-90 (4), P-91 (I), p-n (3), P-II! (I), P-112 (4), P-113 (4), P-I!4 (1), r-ns (1), and P-116 (I). The variations in pathogenicity from the single lesions, single conidium and single cells so far studied are summarized in Fig. S.

Bacterial Blight Disease

Further screening for resistant varieties A total of 607 varieties were further tested for their reaction to bacterial blight in addition to the

Table 3. Differentiation of pathogenic races from single hyphaJ tips of germmating conidia.

Single hyphal tip isolate

Racesdifferentiated International race group 10-9 (16), IA-lO (2) 10-1 (2) Philippine races P-106 (10), P·105(5). P-99 (1), P-l07 (1), P-l09 (1), P-ll0 (1), P·l1l (1) P·23 (6), P·l05 (3), P·l06 (2), P·13 (2). P-2 (1), P-9 (1), P-21(1), P-22(1) P-23 (10). P-8 (1), P-9 (1), P·19 (11. P-21 (1), P-33 (1), P·68 (H. P-l06 (1) p·23 (6), P·l05 (3), P-2 (1), P-l0(1), P·19 (1), P-l06 11 I, P·l07 (1), P·l08 (1)

A-a (20 isolates)

A·b (17 isolates)

10-9 (9), IA-4 (2) IA-l05 (2), IA-l09 (1) IB-41 (lL 10-13 (1) IF-l (1) 10-9 (14). IA-105 (3)

B-a (17 isolates)

s-e

(15 isolates)

10-9 (9), 1C-9(3) IC-l (1 ),10-10 (1) 10-13 (1)

Plant Pathology

85

(% ) L-I (56 Isolates)

) (% l-2-36(25)

25

ffihnl>
L-2(44)

50

Since varietal reaction varies with different strains of the bacterium and was affected by temperature during the conduct of the experiment, the resistant varieties selected from the early tests were further tested against 10 strains of the organism to confirm their resistance (Table 4). Reaction of Japanese varieties to the Philippine strains There are a number of Japanese Varieties that generally are known to be resistant, moderately resistant, and susceptible to bacterial blight. To better understand the interaction of rice varieties and the bacterial strains, some of the Japanese varieties were inoculated with virulent, intermediate, and weak Philippine strains. The results (Table 5) showed that these varieties reacted Similarly to the Philippine strains. They further indicate that most of the Philippine strains are similar to the most virulent strains of Japan (Type lor A). Transmission through seed Rice seeds harvested from diseased fields are often contaminated or infected with the bacterium. In temperate regions, this is not considered important in transmitting the disease to the next planting. In the tropics, however, recent reports have indicated a high percentage of seed contamination and disease transmission to the seedlings. Our preliminary experiments have indicated that it is difficult to isolate and positively identify the bacterium from naturally infested seeds, either by planting the seeds on agar medium or by the dilution plate method, because of the very high frequency of other bacterial contaminations. Ordinary hot-water treatment, or even surfacesterilization with chemicals, failed to remove all the contaminants. About 80 yellowish colonies grown from diseased seeds proved to be nonpathogenic, i.e., were not X oryzqe. A convenient method for studying the problem is by completely sterilizing the seeds with higher water temperature and artificially inoculating them with the bacterial suspension. Preliminary experiments have also indicated that the bacterium cannot be recovered from artificially inoculated seeds 2 to 3 weeks after being kept outdoors during summer. This finding led to the study on the viability of the bacterium at

f
50

L-2-42(25)

50

n
L-3 (50)

Ih.
1-281(41)

50

50

t--T--lL-4 (39)

rL
A-a (20)

50

50

h
25

11
L-5 (24) A-b(l7)

rmn

25

mill
8-Q (11) B-b (15)

25

mm
I-

L-I-43

(25)

50

L-I-49 (25)

25

25

123456169101112~14

rn

l
12345676

Races
Fig. 5. Relative population of pathogenic races originating from single lesions. single conidia and single cells of P. oryzae (L-I to L-5 from single lesions, L-I-43 to L-2-42 and 1-281 from single conidia, A·a to B-b from apical or basal cells of conidia).

7,000 varieties screened in 1965 and 1966. 0 f these, 132 were found to be highly resistant in the field by inoculation of the flagleaf with isolate 8-15. These varie ties will be tested against other strains to confirm their resistance.

86

Plant Pathology

Table 4. R eac ti on of selec ted resistan I varie lie s 10 10 strai ns of X an th om on as orv z ee (Dise ase scales suseepti hie).

to 9, re sistan t. to

Variety Zenith TKM6 Wase Aikoku 3 M.Sungsong Keng Chi-ju Earl II Prol ific Early Prolific Early Prol ific Early Prol ific Lacrosse x Zen ith x Nira P.1. 162319 3 8aifufugoya P.I.209938 Tainan-iku 512 Giza 38 8alilla BJ 1 Sigadis Semora Mangga Tainan 9 221 c/B CII/Br /62/2 OZ·78 OZ·60 00·96 DV·29 OV·52 Dv·2 lck) JC· 70 (ck l

Ace no. 131 237 525 755 1540 1766 1768 1769 1772 2001 2097 2396 2773 2856 2993 3086 3098 3711 4095 4181 6883 7608 8555 8558 9647 8816 8828 8806 9114

81 3 3 0 0 2 2 3 2 4 2 2 1 1 3 3 2 3 3 2 0

86 3 4 0 0 3 3 3 3 3 1 4 1 2 4 5 4 3 3 4 0 5 2 2 2

8u 4 1 0 1 4 3 3 3 3 1 5 1 1 3 4
4

Bj S 3 4

8n 3 2

8·75 3 3 1 2 3 3 3 3 4 2 2 0 2 3 3 2

8·77 3 3 1 3 3 2 3 3 5 2 3

8·57 2 3 0 1 2 2 3 1 4

8·59 3 4 1 3 3 2 3 4 5 3 2 0 1 3 3 4 3 3 3 2 4 4 3 3 3 3 2 7 9

B-23 2 2 0 1 3 2 3 1 2 1 2 0 0 1 3 4 3 2

0
1 3 2 3 4 5 2 2 1 1 3 4 5 4 3 5 1 3 4 2 3 3 3 1

0
1 2 1 3 3 4 1 3 1 1 5

1
2 0 1 2 4 3 4 3 3 0 5 3 2 3 3 3 2 9 9

0
2 3 1 3 2 3 3 0
4

6
1 4 2 3 2 3 3 2 3 3 3 1 9 9

5 2 2 2
4

3
3 3 0
4

2
1 3 1 2 0 1

3
3 2 3 3 3 2 6 9

4 2 3
4

3
3 1 8 9

2 2 8 9

6
9

3 3 3 3 3 3 7 9

3 3 3 3 3 3

2
0 3 7

6
9

Table 5. Reaction of resistant. (R), moderately resistant (M) and susceptible (S) Japanese varieties to virulent (V), intermediate (I), and weak (W) strains of Xentnomones oryzae in the Philippines. (Disease scale 0, 1·9; 0, I are most. resistant, 9 most susceptible. The disease reading was made after 30 days),

Averaqe reaction to Phi Ij ppine strains Variety 8·2 (V) Kidama (R) Koqans-rnaru (R) Norin 27 (R) Norin 35 (R) Norin 121M) Norin 131M) Norin 181M) Sasahigure 1M,) Aichi-asahi (5) Jikkoku (5) Kin-maze (SI Norin 39 ls) Norin 51 (SI Zen ith R (ck)
Tsao-tsusn 1M ck)

8·6 (V) 6 8 6.5 6.5 6 6 5 6.5 6.5 6.5 6 7 3 9 9

8·15 (V)

8·57 (V) 6 9 5 5 7 7 7 6 6 7 4 8 9

8·59 (I) 7 9

B·23 (W) 6 9

6 9

6
6.5 5 6.5 7 7 5 6 6.5 6.5 7 7 4 8

6
5 5

5 5 6 5 7 5 6 4

4 4 3 4 5

JC·70 IS ck)

5 5 7 4 8 9

9
9

4 2 6 5

Plant Pathology

87

100 90 80 70

_~
\•

~-o--o--o--o--o-_-o-

__

\,

, ,, ,\

\\

Cold room (4 Cl
\ \ \ \

100

-l--v-~--*--*-,----l!
"It

90

." r,
\\
I ,

\\

\\

\
\ \

~ 50 <=
(f)

"

..
\

., \

.,
',Air-condiliOned room , (22-24 CI

80

\\> \\I" I,
I,
\ I,

\\

*\ \

36 C 34C 32 C

o
\

"",,-

40 30

"

.,

""

\\ ,\oom
"

20 10

..-, .
"

lemlarclure

(21-23Cl

70 ~60 ~ e; '6

\
,

I0 \

\,

I\

"

'

\I \\ II \.\
\
\

""
"\

\
\\

*
\

29 C 22-24 C

\~\..
\ \

.2: 50

(J)

"

40

\I \,\ \ \ .\ I \\ \ \v\\ II' \., v\\ '.\ \ \ I

\\

\' *\

*
V
\

\\

.',

Days after lncubotion

",
o
•
V

30

,
I

\\

\\

36C\ 20

34C\
\

\\
\

*\\ *
\

32C\
\

\0
\

10 OL_ __ ~ ~ 6

.,

\\

"
\

'.

0\\ 0

\ \, \

Vze c

'*
,22·24C

,,\

\V\, ,II'.
__ ~

'"'''' .\
18

\\

.',.',, ,
""
__ ~'~

\
J_~

,
*

_L_ ... _-~-~~

Fig. 6,A. Longevity of X. oryzae on seeds stored in flasks at different temperature conditions.

12

24

30

36

42

Days otter Incubation Fig. 6,C. Longevity of X. oryzae , on ent temperatures.

rice

hulls

at

differ-

100

I" 90 80 70 60

---v--r-*-*--*-*·-*'·---------, \ I 1\ \* I' I 1 \\ \I , 36 C II

\0

[[IJ

F-ew Col cnl e"!

D No

Growth

t-

"

1\ \I

,\ I' I\ I\ II

I I I

"
1\

\I

II

..
I'

I\ I,

\ \

\..

\
\28 C

~ e;

I' I' II

32 C\

\\ "\ \
I

,I

\ \34 C

.\\
\, \

.,
\
\

I I I ,

28

34 C 32 C

Infected

Leaves

22-24

\22-24 C

\.

\\

Water

Suspension

u
\

\\
36C~\

II

v\ , \I ., • \ \
\
\

\ \11'
I

*),
'

.\ \.

\ II

\\

\\

\ \. \ '\
\

\ \

\ ,\
\

.
\ \ \

\ \ ** *\

'"

""

Agor Siont

'*
o

,\

"

II

'\

L__-L

__

\O-_~ __ 12 18

~~-A~_~

\\ \
30

__

24

36

42

Days after Incubotion Fig. 6,8. Longevity of X. oryzae on rice grains (brown rice) at different temperatures.

12

18

24

so

Days ofter Incubation

Fig. 6,D. Longevity of X. oryzae from different sources at differen t tempera tures.

88

Plant Pathology

Fig. 7. J n fec lion and killing of rice seedlings by ell tting roots in suspension of X oryzae, Seed lings a t left are not infected, the roots are not ell t, even though they are similarly inoculated with tile bacterial suspension.

various temperatures. Infected leaves, rice hull, and brown rice were kept at 24,28,32,34, and 36 e, and some of them at 4 e and at ·30e. The results (Fig. 6A, B, e, D) showed that the bacterium quickly lost its viability at higher ternperatures. At about 32 e the bacterium lost its viability within 30 days. At 4 e, all the inoculated seeds produced bacterial colonies after several months. At ·30 C, the bacterium was readily isola ted from infected leaves after 18 man ths, In the tropics, rice seeds are often exposed to 30 to 32 C or higher for a considerable period, particularly when these were starting to dry up after harvest. It is doubtful that the bacterium on or in the seeds can remain viable up to the next planting if the interval between harvest and the next planting is more than 30 days. Experiments also indicated that seedlings from inoculated seeds and seeds grown on inoculated soil or mineral-cui ture solution are not infected unless the root is inju red or cu t (Fig. 7). The above experiments suggest that in the tropics, infected or contaminated seeds hardly transmit the bacterial blight disease.

Effect of shading and plant age on disease development

Previous results have shown that varieties susceptible to bacterial bligh t have a higher red ucing sugar content than those resistant to the disease. Two preliminary experiments showed that rice plants grown in reduced light (partly shaded) developed smaller lesions than those exposed to full sunlight. The plants in both cases were treated in total darkness for 2 to 3 days before they were ex posed to the two light conditions. In the experiments on plant age in relation to lesion devel opmen t, young seedlings (10 days afte r germination) even of resistant varieties such as Zenith were severely infected or died from needle inoculation. It was inferred that these young seedlings contained large amounts of sugar from the decomposition of starch in seeds. All these experirnen Is suggest that the sugar content of the leaves is closely related to the developmen t of the disease.

Plant Pathology

89

Bacterial Leaf Streak

Varietal resistance

More than 600 indica- and japonica-type varieties and 49 wild rice species and strains have been screened for resistance to the bacterial leaf streak disease using S- J 03, a virulen t strain of the bacterium. The results are summarized in Table 6. Most of the varieties were moderately resistant, and a number of them showed highly resistant reaction. As a group, wild rice is more resistant to the disease than the cultivated forms. The varieties found to be resistant (Scales I and 2) among those tested are listed below for possible use by breeders. The following varieties have reaction Scale 1:
1.

OZ-60 (lRR!

Ace,

No. 8558)

2. Oryzae perennis (100192)

3. Australian spontanea (100944)

4. Australian spontanea (101146) 5. Australian spontanea (101147) 6. Australian spontanea (101148)

The following varieties have reaction Scale 2: J_ Milbuen 5(3) (2 entries) (49) 21. 00-96 (2 entries) (8647) 2. Hsinchu 56 (77) ·22. 00.100 (8649) 3. Ch 242 (87,7114) 2l 00-113 (8657) 4. Bluerose (551) 24. Ctg. 1118 (8712) 5. NOlin 37 (2556) 25. OV-52 (2 entries) (8828) 6. Hashikalmi (3397) 26. DV-29 (2 entries) (8816) 1. Charnock (Aus.)(3685) 27. OV·57 (8831) 8. NP-97 (3700) 28. OV-68 (8833) 9.. Bmt. 53 R3540 29. OV-98 (8845) (4132) 30. OV-150 (8876) 10. Huan-sen-goo (4619) 31. JW-I07 (9129) 11. Pi 4 (6733) 32. Oryzee glaberrima 12. ONJ-142 (8426) (100140) 13. D1-65 (8485) 33. O. glaberrima (100143) 14. OZ-I92 (8518) 34. O. perennis (100211) IS. OZ-179 (8520) 35. O. breviligulata 16. OZ-153 (8538) (l00927) 17. OZ-74 (8556) 36. Australian spontanea 18. OZ-39 (8564) (lOI145) 19. DL-IO (8597) 37. Oryzae rufipogon 20. 00-89 (8644) (101448) As reported earlier, there was a very high correlation between the disease scales based on artificial inoculation and the visual field scales, particularly when virulent strains were used in the inoculation (Table 7). This is understandable since there are many virulent strains in the field. Plant age and disease development Both artificial inoculation and field observations indicated that plants become more resistant to

bacterial bJight at the later growth stage. This increase in resistance was more significant in some varieties than in others, Eight resistant, moderately resistant, .and susceptible rice varieties were inoculated with 8 virulen t and weak isolates of the bacterium at the age of 22, 39 and 57 days, and the sizes of the lesions produced were measured. In all cases, the 57-day-old plants produced significantly smaller lesions, averaging less than one-half the size of those on the younger plants. The difference in the production of lesions between 22- and 39-day-old varieties was not always Significant. As indicated in the foregoing, the resistance may increase further when the plants become older. In another experiment, a few varieties and selections were grown to observe the development of the bacterial blight disease in the field. The disease began to appear 4 to 6 weeks after transplanting, when the leaves of the plants among the hills were in contact with each other. Under favorable conditions in the wet season, the disease became more serious and the susceptible varieties looked completely yellow by the 7th and Sth weeks. In the 9th and 10th weeks, the flag leaves which appeared were, however, almost intact, and since these leaves dominated the field, the plants looked healthy green and remained so until harvest. At the same time, the younger plants of the same varieties were heavily infected. This type of adult plant resistance was particularly significant in such varieties as IR8, IR5 and BPI:76-1. In other varieties, e.g., Peta and C-18, such resistance was less pronounced. The disease is most intense when favorable climatic conditions (high moisture and wind or rain-storm) coincide with the more susceptible age of the plants, The tolerance to the disease at the later growth stage may also be considered as a type of resistance and may be utilized in breeding for resistance.
Relative virulence in the Philippines of the strains

Preliminary inoculation experiments using two susceptible varieties, Pah Leuad 29-8-11 and Pets, indicated marked differences in virulence among the 150 isolates obtained in the Philippines. The results of the experiment in which eight selected strains were inoculated to 16 rice varieties with various degrees of resistance are shown in Fig. 8.

90

Plant Pathology

Table 6. Varietal resistance (fest strain: S· 103.)

to bacterial

leaf streak

of indica and japonica

varieties

and wild rice species and strains.

Nu mber of varieties Resistant Reaction (scale) 2 3 4

per reaction

group Susceptible 6 7 Total no. 9 varieties

Intermediate 5

8
26 9 0 16 51

Indica varieties Japonica varieties Wild species and strains Orhers Total

1 0 5 0 6

30 5 6 0 41

62 15 7 4 88

61 22

8
7 98

110 37 9 13 169

65 24 7 21 117

52 14 7 16 89

5 0 0 1 6

412 126 49 78 665

Table 7. Scales for rating varietal reaction

to bacterial

leaf streak in greenhouse

inoculation

and field visual observation.

Scale

Greenhouse inoculation

Field observation

0 1 2 3 4 5

6
7

8
9

No lesion <1mm 1-2mm 2-5mm 6-10 mm 11-20 mm 21-30mm 31-40 mm 41-60 mm >6Omm

No lesion Sporadic and few lesions Very few lesions per plant Few lesions per plant but all over the field Generally few lesions but some plants heavily infected Many lesions per plant allover the field Many lesions. some leaf tips yellow leaf tips yellow Whole parts of leaves yellow, leaves drying leaves drying

}
}

Field looks green Field looks yellow

Isolates S-103 and 8-107 were the most Virulent, 8-1, 8-64, 8-115, and S-1O1 had intermediate virulence, and S-20 and S-95 were the least virulent (weak strains). The resistant varieties were more resistant to all isolates while the susceptible ones produced large! lesions to all isolates although there were variations within each group. There were no distinct differences in the reaction of the varieties to different isolates. There seemed to be no distinct pathogenic races as in the case of the blast fungus, i.e., those varieties that are resistant to the most virulent strains would be expected to be resistant to other strains.
Pathogenicity patterns

degrees of resistance were selected to study the pathogenicity patterns of the organism or the reaction types of the varieties. Various patterns and reaction types were observed; these may, however, be summarized into four major types (Fig. 9). Type I, represented by isolate 8-78, showed high virulence and a distinct reaction among the resistant, moderate and susceptible varieties; Type II, represented by isolate 8-95, had intermediate virulence; Type III, represented by isolate 8-95, had very weak virulence; and Type IV, represented by 8-94, did not show any marked difference in reaction among the Varieties. These pathogenicity patterns are simiJar to that of the bacterial blight organism.
Bacteriophage population in irrigation water

To better understand the interaction of varieties and strains, 25 isolates of the bacterium of different virulence and 6 rice varieties with various

To obtain information on the fluctuations of the bacterial population in the water of irrigation

Plant Pathology

91

4 2 0 4 2 0 4 u
If)

•
0

5-78 S-93 5-94 5-95 25 Strains overage

.,.
V 4 X

2 0 4 2

a
_j
If) Q)

-e 3 -2
c 0 .iii
<I) _j Vl

a
..c
Q)

0. 0 c
I

a
-£
C

4 2

01

...J

<I> _J

U)

Q) If) Q) If)

i:5

o~~~==~~~~~~
4 2

o~----~----~----~~----~--~6 I 2 3 4 5
Fig. 8. Reaction of 16 rice varieties to 8 isolates of X. translucens f. sp. oryzae. Resistant
1- Charnock (Aust.)
2- N.P. - 97 3- Norin 22 4- Milb ue n 3 S- Milketan 20 6- CI-7 3385 7- Kung-shan-wu-shen-ken S- Un blatu z i Valley Sugar Co.

f-- Resistant Susceptible -4 Varieties

I 2 3 4 5 6 7 8 9 10 II 12 131415 16

Resistant Varieties

Susceptible --------t

Fig. 9. Reactions of resistan t, in terrnediate and susceptible varieties to strains of X translucens f. sp, orvzae.

Susceptible
9- Paikan-tao
10- NP-137

11- Talchung-ti-chio-wu-chien
12- Pe ta

13- Ya-y iu-tsai

.14- Kam-bau-ngan IS- Mamor.iaka 16- Nhta-lO

canals and of fields during rice- growing seasons, the bacterial population was taken by means of bacteriophage counts. Water from three sites was sampled: the large canal leading to the lRRI Experimental Farm, the rainfed reservoir of the same

92

Plant Pathology

Farm, and the drainage- canal leading out of the Farm. The samples were taken at about l-week intervals. The results showed that the water in the drainage canal had the highest bacterial population - 6,000 to 7,000 plagues per ml of water were counted from August to October. The water in the reservoir had the next highest count, while that in the large irrigation canal had the lowest. However, the fluctuation in population was greater in the water of the large canal; occasionally more plagues were detected in this canal than in the reservoir during the early months. A summary of these results into a monthly-average basis from April 1967 to February 1968 is shown in Fig. 10 A. The phage population was also counted in four field plots, two each of a more resistant variety, Hsinchu 56, and a susceptible variety, Pah Leuad 29-8-11. One plot of each variety was inoculated by spraying the bacterial suspension on leaves I month after transplanting. Phage plague counts were made at approximately weekly intervals. The results (Fig. 10,B) showed that: (I) the phage
2000

50000

o
20000 --

Hsinchu 56
Inoculated
Uninoculoled

10000

5000
I

!: 2000 Kl
'1000

I I

I I

I\

/\

'.
\ \ ...

'0
.0

500

200

100

I I

: J , , ,I

I I

, ,
I
I I

20

1000

--

_
_

Lorge IrrigQnon ccocr

R(linred OtaillClge wale, COnOI '~p'\Joir

f---

t:=J

June -

9 14 A_UI,I 1967

500

200

1-

Fig. 10,B. Bacterio-phague counts for X. translucens r. sp, in field wa tel under resistant and susceptible varieties.

oryzae

~IOO

1-

3 g
Ii

50

'0 E

1-

Z
:>

20 -

I-

I-

1-

1-

,--

1-

10

I-

I-

I-

I-

5-

I-

I--

I-

i-

population is much higher in the water in the susceptible varieties than that in the more resistant varieties with or withou t inoculation, (2) the phage population increased rapidly after inoculation for a period of about 5 weeks, after which it declined to a level similar to that of uninocula ted plots, (3) the phage population increased rapidly 4 or 5 weeks afte r transplan ting (wi tb out inocu lation), and (4) the phage population was generally higber in field water than in the water of the large irrigation canal and in tbe water reservoir of tbe Farm. The reason for the dip in July is not known; perhaps it was due to season.
New phages and lysotypes As reported earlier (1964), six phages (Sp-I to Sp-6) and 16 lysotypes of the 84 isolates of the streak organisms were found in the Pbilippines. These phages and the bacterial isolates stored in a deep freezer were tested again for their inter-

1-

Fig. 10,A. Bacteria-phage plague counts for X. translucens f. sp. oryzae in irrigation water, IRRI Experimental Farm (monthly average from weekly counts).

Plant Pathology

93

reactions. Several single-plague reisoiations were made from each of the original phages. These single-plague reisolates differed from the original phages in their reaction to the host bacterial isolates. For instance, a reisola te from Sp-I phage, designated as Sp-I-I, lysed the bacterial isolates 8-20, 8-24, and S-51, all of which were not lysed by the original Sp-I. Also, the bacterial isola tes S-43 and S-81 were lysed by Sp-I but not by Sp-I-I. A new type of phage, therefore, seemed to have appeared. All the original six phages, except Sp-3, produced this new type. Nine such new phages were found and designated as Sp-I-L, Sp·I-2, Sp-2-J, Sp-2-2, Sp-2-3, Sp-4-I, Sp-5-1, Sp-6-1, and Sp-6-2. It appears that the phages are variable and new ones may be produced in the old cultures. Besides those produced by the old cultures, many new phage isolations were made. A number of these were classified as new and differed in their reaction to bacterial strains. Based upon 20 host strains, 18 more new phages were characterized and designated as Sp-7 to Sp-24. Altogether, there are 33 known phages of the streak organism. The details will be presented separately from this report. The ability of the 33 phages to lyse the 20 test bacterial isolates varied from 1 to 14, mostly 5 to 8. The 20 bacterial isolates sensitive to the 33 phages varied from 2 to 28, mostly 10 to 1 5. At least three bacterial isolates in combination, 8-74, 8-73 and 8-77, were required to de tec t the presence of all phages. The study suggested the variability and complexity of the bacterium-phage interaction which are generally used for detecting other phages and which in turn indicate the presence of the bacterium causing the streak disease. Host range None of the 13 species in 10 genera of grarnineous weeds and 10 varieties of sorghum, corn, millet and wheat were infected by artificial inoculation, even after the leaves were pricked with a needle. All the 49 species and strains of wild rice, however, can serve as al ternative hosts. The degree of infection varied greatly among species and strains, as shown below: Oryza spontanea (IRRI Ace, No. 100910) - disease scale 6

(100912)-7 Oryza perennis (100602)-5 (100919)-4 (100606)-5 (100917)-6 (100930)-4 (100918)-6 Oryza breviliguiata (100117)-7 (100945)-4 (100119)-6 (100946P (100120)-4 O. perennis balunga (101156)-3 (100929)-4 " (101173)-3 (100931)-3 Oryza nivara (101512)-7 (100940)-6 (10 1508)-7 (101252)-4 (101507)-5 O'YZl1 glaberrima (100128)-5 (10 1510)-7 (100136)-5 OrYZl1 perennis (100194)-6 (100137)-5 (100195)-4 (100977)-5 " (100196)-6 (100932)-3 (100208)-5 (100152)-3 (100139)-3

O. sativa f spontanea

Others with scales 1 and 2 are listed under varietal resistance.

Tungro Disease
Varietal resistance to tungro disease The search for sources of resistance to the tungro disease was continued, with a total of 3,990 varieties and breeding lines being tested by the previously described mass screening method. Resistant varieties. Of the 596 varieties tested, 44 were classified as resistant while the rest were either intermediate or susceptible to the disease. Seven of these 44 resistant varieties have been reported in previous annual reports. The following 10 varieties and lines showed less than 30 percent infected seedlings in at least two tests:
Basrnati 370 (lRRI Ace. No. 4895) Seratus hari T /36 (5346) Urang-Urangan 89 (6016) Basmati 37 (6448) Tsou-yuen (7315) 22Jb/210/2/1/1/2(7589) 221c/5 311/2/1 (7596) 221/BCII/ll/3 (7606) 268b/Pr!2/2/2 (7624) Padi Kasalle (8261)

Resistance of IR breeding lines. A total of 2,930 selections of 101 IR lines was tested. About II percent of these selections were classified in the resistant group while the rest were either intermediate or susceptible to the disease (Table 8). New vectors In 1967, Nephotettix

apicalis insects collected

94

Plant Pathology

Table 8. Reaction of IR lines to the tungro diseaseas determined by the massscreening test,

No. of selections showing infection 1%) Li ne and parents 0-30 IR 159, Basrnati 370 x TIN)l IR 506, IR8 x IB589A4·18-1f2 x T(N)1) IR 532, IPeta/3 x T(N)I) x TKM 6 IR 580, IR8 x TKM 6 I R 626, I R 8 x (PetalS x Belle Patnal I R 644, I Fl8/2 x IB589A4·18-1f2 x TIN) 1) I R 658, Peta 7 x T(N)1 I FI 661, I RS x (CP231 x SLO·17) x Sigadis I R 644, I R8 x I R 5·273·'·2 I R 665, I R8 x fPeta/S x Belie Patna) I R 748, I R8 x (.Dawnx Pankhari 203) I R 751, I R 8/2 x (Petal5 x Belle Patna) IR 756, IR8/3 x (B589A4·18-1/2 x TIN)1) I R 781 I R812 x (Yukara x TIN)1) I R 822, I R8/2 x Pankhari 203 I R 854, (I R 8 x Pankhari 203) x I R262·43-8·11 IR 874, IR8/2 x (CP· SLOl2 x Nahng Mon s-4) I R 877, I R 8/2 x I Dawn x Pankhari 203) I R 1154. I R8/2 x Zenith Other 81 IR lines Total (101 lines, 2,930 selections) 5 5 3 10 7 2 2 5 2 2 4 4 2 3 125 21 3 61 6 15 326 31 - 60 61 -100 1 25 66 12 4 7 1 7 2 5 7 3 1 5 254 42 2 13 50 397 1.018

5 20 40 35 12

6
16 4 4 17 4 1

6
609 108 1 165 88 150 1.586

from the provinces of Camarines Norte, Camarmes Sur, Laguna, and Nueva Ec ija, all in the Philippines, were found to be capable of transmitting the tungro virus at the rate of 12, 9, 16 and II percent respectively. The results were confirmed in a further study (Table 9). The percentage of active transmitters varied not only among localities where the insects were collected but also among samples collected in the same locality. A higher percentage of active transmitters was obtained when, in addition to the acquisition feeding, a daily reacquisition feeding was provided. For the daily acquisition feeding, the insects was used for inoculation during the daytime and confined on diseased plants for the rest of the day. Some of the "active" individuals were selected and mated through several generations to produce a colony with a high percentage of active transmitters. The. percentage of tungro diseasetransmitting leafhoppers of N apicalis has now reached about 55. The search for other possible leafhopper and planthopper vectors of the tungro disease, besides N impicticeps and N apicalis, was continued. The 10 species of leafhoppers or plant-

hoppers used in the study were identified by Dr. T. Ishihara of Ehime University, Japan, and Dr. M. S. K. Ghauri of the Commonwealth Institute of
Table 9. Percentage of male NephQtettix apicalis insects transmitting the rice tu ngro virus. Insects collected from Bay. Laguna No, of infective insects/no. tested 0/113 1/104 2/100 3/112 Percentageof active transrni tters 0 0.96 2.00 2.68 8.33 14.77 27.17 45.00 2.86 5.00 10.00 10.00 11.40

Los Banos, Laguna

6/72
13188~ 25/92 45/1oo~

Pila,. Laguna

1/35 2/40 3/30 4/40 105/921

Total

• Daily reacquisition feeding in addition acquisition feeding was provided.

to 4-5 days

Plant Pathology

95

Entomology, London, as Balclutha viridis (Mats.), Cicadulina bipunctella (Mats.), Exitianus capicola Stal., Hecalus sp., Macrosteles [ascifrons (Stal.), Nilaparvatalugens (Stal), Nisin atrovenosa Lethierry, Recilia (lnazuma} dorsalis (Motsch.), Tettigella spectra (Dist.) and Sogatella furcifera Horvath. The only successful transmission of the disease from infected to healthy plants was obtained with the leafhopper R. dorsalis. About 6 percent of the field population of this species used transmi tted the disease. Several field collections of this insect transmitted the "S" and "M" strains of tungro, The percentage of "active" transmitters for all collections ranged from 4 to 8. For the other 9 species, tests involving 112 to 664 insects of each species failed to show any disease transmission.
Morphology of N. apicalis and virus transmission General morphology. The morphological similarities and devia tions from typical morphological characteristics of N. apicalis and N. impicticeps have accounted for the difficulties encountered in distinguishing between these two species. However, the morphological features of the insects of both spec ics collec ted in the Philippines and examined at the lnstitu te generally agreed with the characteristics described by Ishihara. But there were deviations in the arrangement of the teeth or spines on the aedeagus as well as in the number of teeth. The teeth are generally arranged in two rows but in some cases, one to four teeth are arranged irregularly or appear somewhat in one row either on one side or on both sides of the au ter area of the two main rows of teeth on the aedeagus. Furthermore, the arrangement of teeth in each of the two main rows may not be symmetrical. Therefore, the number of teeth in each of the two main rows may not be identical. The existence of teeth in addition to those in the two main rows and the asymmetrical arrangement of the teeth in the two main rows have been observed in both N. apiealis (Figs. II and 12) and N. impicticeps (Figs. 13 and 14). The num ber of tee th on the aedeagus varies from 10 to 23, mostly 14 to 17 for N. apicalis and from 4 to 10, mostly 7 to 8 for N. impicticeps.

!] s

Fig. II. Portion of aedeagus of Nephotettix apicalis showing teeth on each side of the two main rows of teeth,

Based on the present results, the following key

is proposed for distinguishing between N. apicalis and N. impicticeps, Aedeagus without elongated paraphyses and hardly constricted below the paraphyses. Style straight. I. Vertex with a submarginal black band. Tegminal spots often present and confluent along the claval suture. Aedeagus with a total of 10 to 23, mostly 14 to 17 teeth . . . . . . . . . . . . . . . . . . . . . . . . .. N. apicalis 2. Vertex without submarginal black band. Tegminal spots present or absent; if present, not confluent to claval suture. Aedeagus with a total of 4 to 10, mostly 7 to 8 teeth ....................... N. impicticeps Attempts have been made to separate the active transrni tte rs of N. apicalis on the basis of their minor morphological features such as location of tegminal spots, length of aedeagus, arrangement of teeth on aedeagus, and number of teeth. The results are summarized as follows: Tegminal spots. The collected samples showed that the tegminal spots mayor may not be confluent to the claval suture. The insects could be grouped into three categories according to the location of the spots. The spots of the majority of the insects examined were confluent to the suture (Table 10). The spots of the other insects were either not confluent to the suture or only one of the two spots was confluent to the suture. However, the frequency distribu tion of the active trans-

96

Plant Pathology

rnitters by the location of the spots seemed to be identical with that of the insects that did not transmi t the virus. Therefore, the Ioca tion of the spots cannot be used as a criterion for determining the transmissive ability of the insect. Length of aedeagus. The length of the aedeagus of the 100 non-active N. apicalis transmitters examined varied from 0.45 to 0.62 rnm with an average of 0.52 mm. The range of the length of the aedeagus of 73 active transmitters was from 0.41 to 0.67 mm with an average of 0.51 rnrn. The frequency distribution of both active and nonactive transmitters by the length of the aedeagus was similar. Since their differences are not statistically Significant, the active and non-active transmitters cannot be separated by the length of their aedeagal shafts. Arrangement of teeth on aedeagus. As mentioned earlier, the arrangement of teeth on the aedeagus may not always be symmetrical. This means that either the number of teeth on each of the two main rows may not be identical or there

Fig. 13. Portion of aedeagus of Nephotettix impicticeps showing one tooth on the left side of the two main lOWS

of teeth.

~J

J
Fig. 12. Portion of aedeagus of Nephotettix apicalis showing asymmetrical arrangement of the teeth. Fig. 14. Portion of aedeagus of Nephoteitix showing asymmetrical arrangement of teeth.

impicticeps

Plant Pathology

97

Table 10. Frequency of tungro Nepbotettix epicetis by location

summarized as follows: Percentage of active transmitters. As mentioned earlier, the percentage of active transmitters of N. Active Nonactive apicalis varied greatly among the samples collected transmitters transmitters Location of spots from different localities. It even varied among the (%) (%) sample s collected from the same locali ty. However, even with the most active colony, the per88.0 Spots con IIuent to 86.8 centage of active transmitters of N. apicalis WaS claval su tu re significantly lower than that of N. impicticeps 2.7 One spot confluent to 3.8 claval suture (Table II). 9.3 Spots not con fl uent to 9.4 As has been known since 1965, the percentage claval suture of infective insects of N. impicticeps increases as Total no. of examined the acquisition feeding period is increased up to 4 75 insects 106 days. In the case of N. apicalis, the percentage of infective insects mayor may not increase propormay be more than two series of teeth on the tionally with the length of the acquisition feeding aedeagus, The 171 insec ts exam ined (98 non -active period. This depends on the activity of the insects. tran sm itters and 73 active transm itters) can be For instance, if a colony with a low percentage of grouped on the basis of symmetrical teeth and active transmitters is used, the percentage of inasymetrical arrangement. The insects in the first fective insects may remain similar even if the group can be further divided according to the length of the acquisition feeding period varies number of teeth per series, which varied from 6 to from I to 4 days. However, the results in Table 11 10; those in the latter group, according to the show that the percentage of infective insects of number of series, which varied from 2 to 4. The both N. apicalis and N. impicticeps was increased majority of the insects, regardless of whether they when, in addition to the acquisition feeding were active or non-active transmitters, had period, the insects were given a daily reacquisition asyrnetrically arranged teeth. There seemed to be feeding period. Therefore, the effects of daiJy no striking difference between the active and non- acquisition feeding on the infectivity of both active transmitters in the arrangement of teeth. species are basically similar. Number of teeth. The number of teeth on the Virus retention period. One of the reasons for aedeagus of the N. apicalis insects examined varied concluding that the tungro virus does not persist in from 10 to 23 with an average of 15.96. About 82 N. impicticeps is that the infective insects do not percen t of the insects examined had 14 to 17 retain their infectivity. The longest virus retention teeth. The number of teeth varied from 10 to 22 period obtained was 5 days. It was found that the with an average of 15.60 for the active trans- tungro virus also does not persist in N. apicalis mitters and from 12 to 23 with an average of (Fig. 15). The longest retention period obtained in 16.23 for the non-active transmitters. The difference was not statistically significant.
virus transmitters 01 male of tegminal spots.

Virus transmissive ability of N. apicalis and N. impicticeps The tungro virus transmissive ability of N. apicalis and N. impicticeps were compared. Transmissive ability refers to the percentage of active transmitters, retention period of infectivity, effect of daily acquisition feeding on both the percentage of active transmitters and the virus retention period in the insect, and effect of length of acquisition feeding on infectivity. The most active N. apicalis colony obtained so far was used. The results arc

Table J I. Comparison of the average percentage of tungro virus transmitters between Nepbotettix apica/is and N.
imp ic ticeps. Percentage Nephotettix sp. Acquisition feeding only of infective insects LSD 5%

With daily reacquisition feeding

N. impicticeps N. apica/is LSD 1%

69.40 1 7.39

83.67 30.13 17.69

11.09 11.67

12.69

98

Plant Pathology