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Rice Breeding: Past, Present and Future: (IRRI)

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J. Genet., Vol. 66, No. 3, December 1987, pp. 195-216. 9 Printed in India.

Rice breeding: Past, present and future

GURDEV S KHUSH*
Department of Plant Breeding, International Rice Research Institute, P.O. Box 933,
Manila, Philippines

I am greatly honored by my selection for this prestigious prize. I express my sincere


appreciation to the Japan Science and Technology Foundation for considering me
worthy of this award.
I have been extremely fortunate to have been able to work with a team of
dedicated agricultural scientists at the International Rice Research Institute (IRRI)
and with dozens of national scientists in rice-growing countries. Such cooperation
has resulted in the development of a series of high yielding varieties with disease
and insect resistance and early maturity that are now planted on millions of
hectares of rice land in Asia, Africa, and Latin America.
Rice was first domesticated in southern China and northeastern I n d i a - p r o b a b l y
i n d e p e n d e n t l y - a b o u t 8,000 years ago. Constant human selection for improved
traits has modified domesticated rice varieties from their wild progenitors so much
that domesticated rices can no longer survive in the wild state. The simple acts of
reaping and sowing, for example, are selective. Primitive humans may not have
known it, but they started the first rice breeding programs when they began to grow
rice plants for their own use. Most primitive farmers have a keen eye and a sensitive
feeling for plants. Millions of rice farmers have applied this keen insight and
sensitivity for thousands of years to select better varieties.
Selection was first practised on the variable and heterogeneous wild and
semi-wild populations, which must have narrowed genetic variability. However,
several mechanisms in primitive agriculture, such as the introduction of varieties
from one region to another and occasional natural crosses enhanced variability for
further selection. Natural crosses between the domesticated crop and the weed
complexes were another source of variability. Tile third source of variability was
the varietal mixtures that primitive agriculturists grew as a protection against dis-
ease epidemics. Occasional intercrosses between component varieties gave still
more variability. This conscious and unconscious selection by humans led to the
developnmnt of over 150,000 varieties grown around the world.
The foundations of modern plant breeding were laid between 1700 and 1900. The
1694 discovery of sex in plants and hybridization beginning in 1719 paved the way
for the artificial creation of variability. During this period, scientists proposed the

*Text of a lecture delivered by the author on 16th April 1987 in Tokyo upon receiving the Japan Prize
for 1987.
195
196 Gurdel, S Khm'h

"cell theory" and discovered the existence of the nucleus, chromosomes and
reduction division to produce gametes and fertilization. The phenonmnon of
sterility was noted in interspecific crosses and the inheritance of characters was
studied. Darwin's books, 77re Origi#z o/" &)ecie,s" by Metros o/" Natural Selectio#l and
The Effects of Cross apid Self Fertilizatio#l it1 the Vegetable Kbzgdom, enlmnced
interest in plant breeding.
The scientific basis of plant breeding has been enhanced tremendously during the
20th century. New breakthroughs have restllted in refinenaents of the two phases of
plant breeding: the evolutionary phase (creation of variability) and the evaluation-
ary phase (selection of superior combinations).
The rediscovery of Mendel's Laws in the first decade of the 20th century
elucidated the mectmnism of segregation of traits and the laws of inheritance. In the
second decade, Morgan and his colleagues mapped Drosophila chromosomes,
thereby explaining the linkage and recombination of traits, vavilov established a
germplasm collection of cultivated species and their wild relatives- the world's first
"gene bank"-in the third decade. This was the forerunner of today's germplasm
banks, which supply tim "genetic building blocks" for modern crop breeding
progralns. Developments in cytogenetics during the fourth decade of this century,
improved our understandilag of the structure and functions of chromosolnes, the
mechanisms of recombination, and the relationships of species. In the fifth decade,
the discovery of colchicine provided scientists a new tool in chromosome
manipulation. The potential of X-rays in generating new variability was the greatest
discovery of the sixth decade. Techniques for inducing gene inutations have
become powerful tools for the plant breeder. Developments in biometrical and
quantitative genetics in the seventh decade'have helped plant breeders formulate
selection strategies and conduct stability analyses.
Farmers themselves were responsible for most rice improvement from the time
of its domestication to about 1900. The best-known examples are the "rono"
varieties such as <'Shinriki" that Japanese farmers selected in the 1890s. The rono
varieties were shorter and therefore responded to nutrient inputs with higher
yields. Rice breeding stations were established in China, India and Japan in the
early 20th century. Rice breeders' initial activities were the purification of existing
varieties through pureline selection. Some crossbreeding, or hybridization, was
done, but varieties of hybrid origin had little impact until the 1950s. In 1949, the
International Rice Commission (IRC) established a breeding program with
headquarters at Cuttack, India, to cross tall tropical "indica" varieties with shorter
'~ from Japan and other regions of Eastern Asia. Most of these
"North-South" combinations failed, but some varieties developed through the
(1P,C), such as Mahsuri and ADT 27, are still popular farm varieties in South and
Southeastern Asia.
In 1956, agricultural scientists in Taiwan, China, released Taichung Native 1-the
first semidwarf rice variety to cross national boundaries in the tropics. TN1 was the
progeny of a cross of Dee-geo-woo-gen, a dwarf rice from China, and a local
variety. Because of its resistance to lodging when fertilized, TN1 was widely grown
in India during the mid 1960s.
Despite the impact of varieties such as TN1, Mahsuri, and ADT 27, rice farmers
in tropical and subtropical Asia grew hundreds of thousands of unimproved or
semi-improved varieties even in the early 1960s, and few had l~een touched by
Rice breeding: Past, present and .fi~ture 197

modern agricultural science. These varieties were tall and weak-stemmed, and
were late maturing. Most were sensitive to photoperiod or daylength, so they could
only be grown in limited geographical regions. Such varieties, if resistant to one or
two specific local diseases and insects, were invariably susceptible to those from
neighbouring areas. Average yields in most tropical rice-growing nations were
from 1 to 2 tons per hectare.
In the late 1950s and early 1960s, populations ira these countries were increasfi!g
faster than food production. Because of these trends, several authorities such as
Paddock and Paddock (11967) and Borgstrom (1972) predicted large-scale famines
in the late 1970s. However, recent advances in wheat and rice breeding have
produced varieties with a yield potential 2-3 times higher than that of the
traditional varieties. These new varieties respond better to modern agronomic
practices. Wide-scale adoption of these varieties and proper agronomic practices
have led to major increases in foocl procluction World rice procluction increased
8 2 % - f r o m 257 million tons ira 1965 to 468 million tons in 11985. World wheat
production increased 89% from 267 million tons in 1965 to 506 million tons ira 1985.
As a result of the Green Revolution ushered in by improved rice and wheat
varieties we now have 450 million tons of food grain reserves worldwide. Thus,
instead of food scarcity and famines we have problems associated with overproduc-
tion, such as storage of surplus grain and crashing of prices.
Despite this overproduction, people in certain regiOns cannot afford adequate
amounts of food. The Green Revolution technology has bypassed certain regions.
The world population is likely to double in the next 40 years and we nmst double
our food production in the same period.
I shall now attempt to review the major advances that have taken place in rice
breeding leading to the Green Revolution in rice farming, and the challenges which
lie ahead.

Recent achievements in rice breeding

The International Rice Research Institute (IRRI) was established ira 1960 to apply
science to agriculture to increase the production of rice, which forms more than half
of the total food consumed by one out of three persons on earth. Ira 1962, IRRI
scientists crossed Dee-geo-woo-gen, the same Chinese variety which had given TN 1
its semidwarf plant stature, with Peta, a vigorous variety from Indonesia. In late
1966, IRRI released the variety IR8 from this cross. Because of its superior yield
potential, IR8 was widely accepted. Other improved varieties were developed ira
succession.
_ The, improved rice varieties which are now planted on 60% of the world's
riceland have many desirable features which were not present ira the pre-Green
Revolution varieties. These features are 1) high yield potential, 2) short growth
duration, 3) multiple disease and insect resistance, 4) superior grain quality, 5)
tolerance for problem soils.

Yield potential

Rice varieties cultivated by farmers in the tropics and subtropics in the pre 1I),8 era
were tall and leafy with weak stems, and had a harvest index (ratio of chy grain
198 Gm'dev S Khush

weight to total dry matter) of 0.3. When nitrogenous :fertilizer was applied at rates
exceeding 40 kg/ha, many traditional varieties tillered profusely, grew excessively
tall, lodged early, and yielded less than they would with lower fertilizer inputs. To
increase the yield potential of tropical rice, it was necessary to improve the harvest
index and increase lodging resistance and nitrogen responsiveness. This was
accomplished by reducing the plant stature through incorporation of a recessive
gene for short stature from a Chinese variety Dee-geo-woo-gen. IR8 also had a
combination of other desirable features such as heavy tillering, dark green and
erect leaves, and sturdy stems. It responded to nitrogen fertilizer much better than
traditional varieties such as Peta (figure 1). It had a harvest index of 0.5 and it
doubled the yield potential of tropical rice. It was photoperiod-insensitive and had
a growth duration of 130 days. Ttms, in the tropics it could be planted any time of
the year.
The IR8 plant type concept was so convincing and revolutionary that all the rice
improvement programs immecliately initiated crossing programs to d e v e l o p
short-statured varieties. Since then, more than 500 short-statured varieties have
been developed by IRRI and national rice improvement programs, which are now
planted on most of the irrigated and favorable rainfed lowland areas. During the
20-year period 1966-86, short-statured rice varieties were adopted on 60% of the
riceland. Perhaps no other innovation in agriculture has had this wide acceptance.

Short growth duration

Most traditional varieties in tropical and subtropical countries mature in 160-170


days and many are photoperiod-sensitive. These were suitable for growing one crop
of rice a year during the rainy season but not for multiple cropping systems. IR8
and subsequent varieties such as IR20 and IR26 mature in about 130 days.
However, if the farmers grow 130-day-duration varieties, it is not possible to grow

9,000 / ~ IR B

o 8,000

133
. - 6,000

'c~
79. z,,000 " `%`%
,%

"~"~'Peto
2,000 I ..............................................................................................................................
0 - 30 60 90 120
Nitrogen opptied,kg/ha
Figure I. Nitrogen response of Peta and 1R8 during 1966 dry season at IRRI.
Rice breeding: Past, present and fitture 199

another crop after rice or a second crop of rice in one rainy season. Therefore,
major emphasis was placed on developing improved varieties with shorter growth
durations. IR28 and IR30 developed in 1974 and IR36 developed in 1.976 mature in
110 days. The growth duration was further reduced to 105 in IRS0 and 100 days in
IR58. However, yield is the primary consideration in the varietal development
program. During the selection process, only those short-duration lines with yield
potential that matched that of medium-duration varieties were saved. The key to
the success of.this program was the selection of genotypes with rapid vegetative
vigor at earlier growth stages (Khush 1987). Because of higher growth rates at
earlier stages, the short-duration varieties such as IR36, IR50, or IR64 are able to
produce approximately the same total biomass in 85-90 days as the medium-
duration varieties do in 115-120 days. Moreover, the harvest index of short-
duration varieties is slightly better than that of the medium-duration varieties.
Under most situations, the yields of early and mediurn growth duration varieties
are similar.
However, because the short-duration varieties produce the same amount of grain
in fewer days than medium-duration varieties, their productivity per day is much
higher. In replicated yield trials at IRRI, IR64 produced 88 kg/day during the 1.984
dry season and 54 kg/day in the wet season (table 1). IR42 yielded 68 kg/day in the
dry season and 39 kg/day in the wet season.
Short-duration varieties are excellent for input economy. Because they grow
rapidly during the vegetative period and are thus more competitive with weeds,
weed control costs are reduced. Pesticide requirements are also minimized. And
because the field duration of these varieties is 25-30 days shorter, they utilize less
irrigation water, thus lowering production costs.
The availability of short-duration varieties has led to major changes in cropping
patterns in Asia. In the Philippines, many farmers grow an upland crop either

Table 1. Yield of promising short-duration lines and wu'ieties. IRRI, 1984 dry and wet
seasons.

1984 DS 1984 WS
Growth
Selection duration Total Yield Total Yield
(days) yield per day yield per day
(t/ha) (kg) (t/ha) (kg)

IR 19743-46-2-3-3-2 98 5.6 73 4.2 54


IR 29658-69-2-1-2 107 6.8 79 4.8 56
IR 29658-94-2-1-3 107 7.4 86 4.9 57
IR 29725-22-3-3-3 108 7.1 82 3.8 44
IR 31802-48-2-2-2 106 6.7 79 3.6 42
IR 31851-63-l-2-3-2 1106 7.6 89 3.6 42
IR 31868-64-2-3-3-3 110 7.1 80 4.2 47
IR 323117-107-3-2-2 110 7.3 82 4. l 46
IR 32429-47-3-2-2 105 7.6 90 5-7 68
IR 58 100 6.5 82 4.6 58
IR 36 110 7.2 81 4.4 49
IR 64 110 7.9 88 4.8 54
IR 42 (check} 135 7.8 68 4.5 39
200 Gurdev S Khush

before or after rice under rainfecl conditions. In some areas, two crops of rice are
regularly grown during the rainy season (figure 2). In Iloilo province, for example,
70% of the rainfed area is clouble-cropped with rice. Farmers grow three crops in
irrigated areas. In Indonesia, where IR36 and other short-duration varieties have
been widely adopted, the area under rice double cropping has increased rapidly.
These changing cropping patterns have resulted in increased food supplies, higher
food security, and more opportunities for on-farm employment for Asian farmers.
Many national rice !mt?rovement programs have also developed short-duration
varieties. Ratna, Cauvery, and TKM9 in India, BG34-7 aim BG367-7 in Sri
Lanka, ancl Chandina in Bangladesh are well-known.

Multiple dis'ease and insect resistance

Major changes have occurred in the varietal composition and cultural practices for
rice during the post-IR8 era. A relatively small number of improved varieties have
literally replaced thousands of traditional cultivars, thereby reducing the genetic
variability of the crop. Farmers have started using improved cultural practices, such
as application of more fertilizers and establishment of higher plant populations per.
unit area. Development of irrigation facilities and availability of short-duration,
photoperiod-insensitive varieties have enabled the farmers in tropical Asia to grow
successive crops of rice throughout the year.
Reduced genetic variability, improved cultural practices and continuous crop-
ping with rice have increased the genetic vulnerability of the crop. Chemical control
o1: diseases and insects for prolonged periods in tropical climates is very expensive
and impractical. The use of host-resistance for disease and insect control is the
logical approach to overcome these production constraints. Therefore, IRRI'S rice
improvement program has placed major emphasis on developing germplasm with

60 / ~ ~ Seed bed

~E
O

B
+
ff
/ \
rice transplanted .
o
/ I~,-,!.lI:;".+,i+.l ........... ] 1, uplond crop 1 new pottern 3
r
/ }2!.(ii:]ricetransplooted] -1
C / ~ " ~ " jnewpattern2
o 20

.'I ....;..... . !:: /!:~"


rice lbroaacast
............. ~x,J
P,~
new pattern 1
tic; transptanted ( otd'~paYtern++~
+++/
0 ......... J l ..................I..... I
+ l I I. I I I ....l....... '

A M J J A S 0 N D J F M
Figure 2. Scheduleof old and new cropping patterns in the PhilipphmssuperinllX~scdon
rainfall pattern.
Rice breeding." Past, present atM fimr 201

multiple resistance to major diseases and insects. Many national programs have
similarly given priority to developing varieties with multiple disease and insect
resistance.
Five diseases (blast, bacterial blight, sheath blight, tungro, grassy stunt) and four
insects (brown planthopper, green leafhopper, stem borers, and gall midge) are a
common occurrence in most countries in tropical and subtropical Asia. Breeding
programs are focusing attention on developing germplasm with multiple resistance
to these major diseases and insects.
For a host-resistance program to succeed, there must be donors for resistance.
Thus, we need a large germplasm collection to identify donors. We need techniques
for evaluating the germplasm to identify the donors and to screen the segregating
populations. Finally, we need interdisciplinary cooperation between breeders,
plant pathologists, and entomologists for rapid progress.
Fortunately, IRRI and national rice improvement programs !aave assembled
excellent collections of rice gerlnplasm. Pathologists and entomologists have
developed screening techniques and many donors for resistance have been
identified (Khush 1977). Utilizing these donors, we have developed improved
varieties with resistance to as many as four diseases and four insects. The IRRI
varieties IR5, IR8, IR22, and IR24 were susceptible to most of the diseases and
insects. The first variety with multiple resistance was IR26, released in i973. Since
then many varieties with multiple resistance have been developed at IRRI and by
the national programs. Table 2 shows the disease and insect ratings of the IR
varieties.
Large-scale adoption of varieties with multiple resistance has helped stabilize
world rice production. The value of multiple resistance in imparting yield stability is
illustrated in figure 3. As this figure shows, the yield of susceptible IR8 fluctuates
from year to year. If there is a disease or an insect attack, the yield is drastically
reduced. However, if the disease or insect incidence is low, the yield is high. On the
other hand, varieties with multiple resistance such as IR36 and IR42 show only
minor fluctuations in yield from year to year and thus have greater yield stabilityl

10000 I[ = = IR 8
o olR36
IR 4 2
8 000

~.4000
~~176176
I1 Figure 3. Yields of IR8 IR36,
and IR42. Yields of multiple re-
sistant IR36 and IR 42 show little
year-to-year variation; yield of
susceptible IR8 fluctua:tes
2 000 I I I I I I I t I widely. Dry season replicated
1973 75 77 79 81 83 85 87 yield trials at IRI~,I.
YeQr
202 Gurdev S Khush

Tal}le 2. D i s e a s e and insect r e s i s t a n c e o f v a r i e t i e s n a m e d by IRRI ( I R 5 to I R 3 4 ) a n d of IRRI


lines n a m e d as wlrieties by the Philippine G o v e r n m e n t ( I P , 3 6 - I R 6 6 ) .

Reaction"

IR Bacterial Grassy BPH b Green Stem Gall


wlriety Blast blight stunt Tungro biotypes leaf- borer midge
hopper
1 2 3

IR 5 MP, S S S S S S R MS S
IR 8 s S S S S S S R S S
I R 20 MR R S IvlR S S S R MR S
I R 22 S R S S S S S S S S
I R 24 S S S S S S S R S S
I R 26 MP, P, MR MR P, S R P, MR S
IP, 28 R R R P, P, S R R MP, S
I R 29 R R R R R S R R MR S
I R 30 MS P, P, IvlR R S R R MR S
I R 32 MP, R R MR R S R R MR S
I R 34 R P, R P, P, S P, R MR S
IP, 36 P, R R R R R S R MR R
I R 38 P, R R R P, R S P, MP, R
IR40 P, R P, R R R S R MR P,
I R 42 R R R R R R S R MR R
IP, 44 R R S R R R S R MR S
I R 46 R R S MR R S R MP, MR S
IR 48 R R R. R R R S R MP, S
IR 50 MS P, R R R R S R MR S
IR 52 MR P, R R R R S P, MP, -
IR 54 MR R R R R R S R MR -
IR 56 R R R R P, R R R MR -
IR 58 R R R R R R S R MR -
IR 60 P. R R R R R P, R MR -
IR 62 MP, R P. R P, R R R MR -
IR 64 MP, R P, P, P, MR R R MR -
IR 65 R R R R R R S R MS -
IR 66 MR R R R R P, P, P, MR -

" R = resistant, M R = m o d e r a t e l y r e s i s t a n t , S = susceptible, MS = m o d e r a t e l y suscep-


tible; - = reaction not k n o w n . R e a c t i o n s w e r e b a s e d on tests c o n d u c t e d in the
Philippines for all diseases a n d insects e x c e p t for gall m i d g e w h i c h w a s c o n d u c t e d in
India. bBPH = brown planthopper.

Grain quality

Grain quality in rice is dependent on milling recovery, grain size, shape, and
appearance, and cooking characteristics. Higher milling recovery is a universal
requirement and, to some extent, determined by the size, shape, and amount of
chalkiness.in the grain. Most consumers in the tropics and subtropics prefer long or
medium long and slender translucent grains. Grain chalkiness (as in IR8) causes
low milling recovery and poor consumer acceptance.
Rice breeding: Past, present and fitture 203

Tile cooking quality is determined largely by tile amylose content and


gelatinization temperature. In the tropics and subtropics, varieties with intermedi-
ate amylose content and intermediate gelatinization temperature are preferred. In
tile temperate areas of China, Korea, and Japan, however, rices of low amylose
content and low gelatinization temperature are preferred.
hnproved varieties like IR8 and 1R5 have poor grain quality. They have bold,
chalky grains which give very poor lnilling yields. The grains cook dry because of
high amylose content, and have poor consulner acceptance. Improvement of
milling recovery and grain appearance received immediate attention in the
breeding program at IRP,I. All the IR varieties released after IR5 and IR8 have
slender and translucent grains and have very good milling recovery. However,
improvements in the cooking quality were slow because all the donors for disease
and insect resistance used in the hybridization program had high amylose content
and low gelatinization temperature. Tile first improved variety with intermediate
amylose content was IR48, but it has low gelatinization telnperature. Similarly,
several IR varieties such as IR20, IR32, IR36, and IR46, have intermediate
gelatinization temperature but high amylose content. The first IR variety with a
desirable combination of intermediate amylose content and intermediate gelati-
nization temperature is IR64 (table 3). It has been widely accepted as a high quality
rice in Philippines, Indonesia, and Vietnam. Its grain quality is considered superior
to that of the high quality traditional varieties such as C4-63, Azucena, and Milfore.
Another important quality characteristic by which premium quality rices are
known is their pleasant aroma. None of the improved varieties are aromatic.
However, several improved breeding lines with aromatic properties have b e e n
developed and are being evaluated in replicated yield trials. It is hoped that we will
soon have improved aromatic varieties.

Tolerance to problem soils

Millions of hectares of lands suitable for growing rice remain unplanted because of
severe nutritional deficiencies and toxicities. Even well-managed ricelands suffer
from mild nutritional deficiencies or toxicities. For example, Zn deficiency in rice
soils is becoming a common concern in many countries. A vast majority of rice soils
have varying levels of salinity or alkalinity. Several improved varieties have been
developed which have moderate to high levels of tolerance for several nutritional
deficiencies and toxicities. IR36, for example, has tolerance for salinity, alkalinity,
peatiness, and iron and boron toxicities. It is also tolerant of Zn deficiency (IRRI
1982). IR42 similarly has a broad spectrum of tolerance for many soil problems
(table 4). Varieties tolerant of these deficiencies and toxicities have a more stable
performance and do well across several locations. As shown in figure 4, IR36 and
IR42 consistently yield better than IR8 at all levels of fertility. This superior
performance is largely attributed to the broad-spectrum tolerance of IR36 and IR42
for problem soils.

Combination of filvorable traits

For large-scale adoption, a variety must have a favorable combination of traits.


High yield or early maturity alone, for example, is not enough. Therefore, we have
204 Gurdev S Khush

Table 3. Main characteristics of IR varieties.

Variety Environment Growth t leight Amylose Gelatinization Grain size and


suited l'or duration (cm) content temperature shape
(days)

IR 5 Rainfed 140 130 High Intermediate Meclium, medium


IR 8 Irrigated 130 100 High Low Long
IR 20 Irrigated 125 110 High hltermecliate Medium, meditun
IR 22 Irrigated 125 90 High Low Long, slender
IR 24 Irrigated 120 90 Low Low Long, slender
IR 26 Irrigated 130 100 High Low Medium, medium
IR 28 Irrigated 105 100 High Low Long, slender
IR 29 Irrigated 115 100 Glutinous Low Long, slender
IR 3(1 Irrigated 110 100 High Intermediate Medium, medium
IR 32 Irrigated 140 105 High Intermediate Long, slender
IR 34 Irrigated 130 125 High Low Long, slender
IR 36 Irrigated 110 85 High Intermediate Long, slender
IR 38 Irrigated 125 100 High Intermediate Long, slender
IR 40 Irrigated 120 100 High Intermediate Medium, medium
IR 42 Irrigated 135 110 High Low Medium, medium
IR 43 Upland 125 110 Low Low Long, slender
IR 44 Irrigated 130 110 High Low Long, slender
IR 45 Upland t25 100 High Intermediate Long, slender
IR 46 Rainfed 130 110 High Intermediate Long, slender
IR 48 Irrigated 140 120 Inter- Low Long, slender
mediate
IR 50 Irrigated 1(15 90 High Intermediate Long, slender
IR 52 Rainfed 115 95 High Low Long, slender
IR 54 Irrigated 120 95 High Low Long, slender
IR 56 Irrigated 110 90 High Low Long, slender
IR 58 Irrigated 100 80 High Low Medium, medium
IR 60 Irrigated 108 95 High Low Long, slender
IR 62 Irrigated 115 110 High Intermediate Medium, medium
IR 64 hrigated 115 105 Inter- Intermediate Long, slender
mediate
IR 65 Irrigated 115 105 Glutinous Low Long, slender
IR 66 Irrigated 110 100 High Intermediate Long, slender

endeavored to combine most of the desirable traits into the same variety. IR36
released in 1976 was the first product of such an effort. It has excellent yield
potential and a harvest index of 0.55. It matures in 110 days, and has excellent long,
slender grains, multiple resistance to major diseases and insects, and tolerance for
several nutritional toxicities and deficiencies. Because of these desirable attributes,
it was accepted widely and became the most widely planted variety of rice or any
other crop the world has known. During the early 1980s it was planted in 11 million
hectares of riceland all over the world. It still is the most widely planted variety of
rice in the world. It has been estimated that farmers who planted IR36 between
1980 and 1.984 harvested 1 billion dollars worth of extra rice annually. An External
Review Panel consisting of 12 eminent scientists examined the work of IRRI in
1981-82 and pointed out, "The impact of IR36 alone would more than justify the
investment in IRRI since its establishment in 1960."
IR50, released in t980, is another variety with a desirable combination of traits.
It matures 5-7 days earlier and yields 10% more than IR36. However, its
Rice breeding: Past, present and future 205

Table 4. Reaction of some IR w~rieties to adverse soil conditions a.

Toxicity Deficiency
Variety
Salt Alkali Peat Iron Boron Phosphorous Zinc

IR 5 4 7 0 6 4 5 5
IR 8 3 6 5 7 4 4 4
IR 20 5 7 4 2 4 1 3
IR 28 7 5 6 4 4 3 5
IR 36 3 3 3 3 3 7 2
IR 42 3 4 5 3 2 3 4
IR 48 4 7 5 6 0 5 5
IR 64 3 3 4 5 4 4 4

"On a scale of 0-9: 0 - n o information; 1-almost normal plant; 9 - a l m o s t dead or


dead plant.

acceptance has been hampered because its resistance to blast "broke down" in
certain areas. IR64, released in 1985, combines all the desirable traits of IR36 but
has superior grain quality because of its intermediate ainylose content and
intermediate gelatinization temperature. It outyields IR36 by 20% and has
horizontal resistance to blast. It is accepted widely in many countries.

Impact of modern varieties

To date, 30 improved IR varieties have been released by IRRI (IR5-IR34) and the
Philippine Seedboard (IR36-IR66) . In addition, 154 varieties have been selected
and released by the national rice improvement programs from IRRI-bred materials.
About 350 improved rice varieties have been developed by the national rice

0,I
,>,,

o
t92
Fignre 4. Grain yield response of 4 rices to dif-
ferent levels of nitrogen. Data are averages for IRRI
and the three expetiment stations of the Philippine
I I I I I Bureau of Phmt Industry (Maligaya, Bicol, and
0 30 60 90 120 Visayas), 1976-1985 wet seasons. (S K De Datta,
N applied,kg/ha IP,l',l-Agronomy Department, unpublished).
206 Gurdev S Khush

'Fable 5. Total area planted to rice and to improved wuieties in


selected Asian countries, 1986.

Country Total area Area planted to improved


(million ha) varieties %

Bangladesh 10.1 28
Burma 4-6 53
China 33-2 98
India 41.2 53
Indonesia 9.8 77
Korea (South)* 1-2 95
Mrdaysia 0.7 70
Nepal 1.3 30
Pakistan 2-0 51
Philippines 3.2 90
Thailand 9.6 15
Vietnam 5.7 51
Sri Lanka 0.9 91
* Includes firea planted to improvedwtrieties of indica/japonica origin
(25%) and improved japonica varieties (70%).

improvement programs. The improved rice varieties are now planted in 60% of the
world's riceland. In China, Korea, Philippines, and Sri Lanka, more than 90% of
the area is now planted to improved varieties (table 5). In Indonesia, India,
Pakistan, Burma~ Malaysia, and Vietnam more than 50% of the area is planted
to such varieties. Because of large-scale adoption of improved varieties and
associated management practices, rice production has dramatically increased in
most of the major rice-growing countries (table 6).
The Asian rice belt, which used to be a rice-deficit area, has now more than 8
million tons of exportable surplus. Indonesia, which used to import up to 2 million
tons of rice until the early 1980s, became self-sufficient in 1984 and had an
exportable surplus in 1985 and 1986. Indonesian rice production increased

50

40

0
~, 30
E
=-
o
V)
zl
2C
s
Q_

:::::::::::::::::::::::::::::::::::::::::::::::::::::::::::•••.:.•.•.•.••••:.:••::••.•••::.•.:::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::

n L ~!~}~!ii~!i!~!!i~!~!i!;i!~!i!!i!!!!!i!~!!~;i!~!~!~:~i!i!~:!ii~!i!~!~i~!~!!i~:!!!i!i:~:~i!~:!!~!i!i!~!:~!:~!~i~!i!~!~ii!~!~
~1965 67 69 71 73 75 77 79 81 83 85 Figure 5. hnpact of improved tech-
Year nology on Indonesian rice production:
Rice breeding." Past, present and fimtre 207

Table 6. Rice production increases in selected Asian


countrms, 1965-85.

Rice production % increase


(million tons) in 1985 dver
Country 1965
1965 1985

Bangladesh 15.7 22.5 143.3


Burma 8.0 14.5 181.2
China 92.(t 17l .3 186.1
India 45.9 96.3 2(/9.8
Indonesia 12.9 39.1 302.3
Korea (South) 4.8 7.8 162.2
Malaysia 1.2 1.8 150.0
Nepal 2.2 2.8 127.2
Pakistan 1.9 5.0 263. I
Philippines 4.0 9.1 227.5
Thaihmd 11.1 19.5 175.6
Vietnam 9.8 15.8 16l. i
Sri Lanka 0.8 2.3 287.5

threefold in a 20-year p e r i o d - f r o m 12.9 million tons in 1965 to 39.1 million tons in


1985 (figure 5). Rice yields in Indonesia increased f r o m 1,76 t/ha in 1965 to 4.0 t/ha
in 1985. In India, rice production d o u b l e d in a 20-year period f r o m 45.9 million tons
in 1965 to 96.3 million tons in 1985 (figure 6). In some states of India, rice
production dramatically increased. I n P u n j a b , for example, the average yield
increased f r o m 1.6 t/ha in 1.965 to 5.0 t/ha in 1985. T h e area planted increased f r o m
0.29 million ha in 1965 to 1.7 million ha in 1985. Rice production increased f r o m
0.34 million tons in 1965 tO 8.6 million tons in 1985, an increase of 24 times in a
20-year period (figure 7). Rice p r o d u c t i o n in P u n j a b increased 14% per year. T h e r e
is no parallel example of increased f o o d p r o d u c t i o n a n y w h e r e in the world.
Similar increases occurred in t h e Philippines w h e r e rice p r o d u c t i o n increased
from 4.0 million tons in 1.965 to 9.1 millions in 1 9 8 5 - 2 2 7 - 5 % increase in a 20-year

I00

83
I/I

2
o 60
c
.o
u 40

2
o~ 20

0 =========================================================================================================================================================
1965 67 69 71 73 75 77 79 81 83 85 Figure 6. Impact of improved ted>
Year nology on Indian rice production.
208 Gurdev S Khush

~1 Area (million ha)


7 ~:.i:.ii!!Yield (tons/ha)
88 Production (million tons)

5
iiiiiiil ........

2
i'i'i'i
+:+:
:::::::: :~:~:~:~
]
+:+:
:+:+ +:+:
:.:+:.
ii!ilili 9""-"
i~i!~
"'" ~i~!ii~:888 ~ ~ ~ J :i:i:~:~
~:~:~:~:
::::::::
0
1950-51 1955-56 1960-61 1965-66 1970-71 1975-76 1980-81 1985-86
Figure 7. A r e a planted to rice, yield and production of paddy in P u n j a b , 1950-1985.

period (figure 8). This increase is entirely due to increased per hectare productivity
as the cultivated area in 1965 and 1985 was about the same.
The major impact of the increased rice production has been the substantial
decline in rice prices particularly from the late 1970s to the present. As shown in
figure 9, the index of paddy production accelerated faster than the index of real
prices for Burma, India, Indonesia, Philippines, Sri Lanka, and Thailand
(Gonzales and Umali 1985). The real domestic prices in these countries declined
steadily. Lower real prices clearly benefit the consumers- lower prices have income
effects, that is the consumers can gain increasing purchasing power, and therefore,
indirectly imply an improvement in their nutritional status.
The price of rice on the international market has similarly declined (figure 10),
which is another measure of the success of new rice technotogy spearheaded by the
improved varieties.
10

!iiiii!i!i!ii!iijiJi"!ii!iilii Year
Figure
nology on
8. hnpact
Philippine
of improved tech-
rice production.
Rice breeding." Past, present and fitture 209

~ ~ o
6
8
e~l)

E
x

._=
.s o ca
'r-

~ _ aS 8

e w o: E
"V3

I I I I I I I I I Ii~

_ o

ee
,_
o
g.
._=
.g
e-.
to
o

o
LO
0 0 0 0 0 0 0 0 0 0 0 0 O~
"~ to o ~o o LO 0 ~0 LO 0 tO 0 tO
-~ ~0 ~0 Od OJ ~ OJ OJ
210 Gurdev S Khlxsh
10

c
o

~-~5
U3

o
o

0 I I I I I I I I ' I I i I I I I i I I I I I I I
1960 '65 '70 '75 '80 '8! Figure 10. Rice export price at
Yeor Bnngkol~ (deflated)

Future challenges in rice breeding

The biggest challenge that rice scientists face today is how to continuously
generate technology that will lead to increased production commensurate with
increases in world population. It took perhaps one to two million years for the
human population to grow to one billion, which it reached in about 1830. It
required only 100 years to reach 2 billion by 1930 and the three billion figure was
reached in 1960 in only 30 years. The 4th billion was added in 15 years (by 1975)
and the 5th billion in 12 years (by 1987), Although the percentage of annual
increase worldwide has fallen from 2.0 to 1.7% in the last 5 years, the annual
increase in human numbers continues to rise at a frightening rate because of a
larger population base. Each year there are 94 million new mouths to feed. As
shown in figure 11, the world population is likely to reach 8 billion by year 2016 a n d

Billions YeGFs
2016 8 1
.8
2008
,10
1998
.6t 11
The wortd's population growth
past end projected
1987
t ,12

197/t-,
1969111
196~
,15

,30

1~ .100

From the
'beginning
I I I I i I i i I___L___L_--I 0
AD 400 800 1200 1600 2000
Years

Figure II. Wor!d's popuhition growth, past and projected,


Rice breeding: Past, present and future 21]

10 billion by 2030. It should be remembered that the population in the


rice-consmning countries is increasing faster than in the rest of the world. The
number of rice eaters will probably double during the next 30-35 years: Thus, rice
production will have to be doubled by the year 2020. It has been estimated that the
demand for rice will exceed production by the end of this century (IFPRI 1977).
Rice-growing environments are classified into five major categories (Khush
1984). Irrigated rice accounts for about 55% of the total area. Another 5% is
favorable rainfed lowland where improved varieties and technology have been
adopted. Abot~t 80% of the world rice production comes from 60% of these
favorable rice-growing areas. However, very few varieties and little new technology
have been developed for the unfavorable (rainfed lowland, upland, deepwater, and
tidal wetland) environments. The challenges of rice improvement now therefore
are twofold:
(1) For the favorable environments, to develop rice varieties with higher yield
potential, yield stability, superior cooking quality and shorter growth duration.
(2) For the unfavorable environments, to develop improved varieties with higher
productivity, yield stability, and with tolerance to environmental stresses.

Varietal development for favorable environments

Yield: Since the initial breakthrough in yield potential when IR8 was developed,
there has been only marginal increase in yield potential, although per day
productivity has been increased in the short-duration varieties. Our major
challenge is to develop varieties with a quantum jump in yield over that of IR8.
Yield is a function of total dry matter and harvest index. Therefore, yield can be
increased by increasing either the total dry matter or harvest index, or both. Rice
physiologists believe that physical environment is not limiting for increasing the
rice yield beyond current levels. Rice varieties with higher yield potential must re-
tain the "good plant-type traits" namely, short stiff (lodging-resistant) culms, erect
leaves, and high tillering ability. In addition, incorporating the following varietal
traits should help increase rice yield per crop.
(1) Increased harvest inclex:
a. Increased sink size:
(i) Large spikelet number per shoot with suitable spikelet size;
(ii) greater partition of assimilates to spikelet formation.
b. Increased spikelet filling:
(i) manipulation of canopy senescence;
(ii) higher quality grain percentage;
(iii) maintenance of healthy root system;
(iv) increased loclging resistance.
(2) Increased biomass production:
a. Establishment of desirable canopy structure:
(i) rapid leaf area devlopment;
(ii) rapid nutrient uptake.
b. Reduced carbon coilsumption.
(3) Increased canopy photosynthesis during entire growth period.
The yield increase in rice varieties developed so far through conventional
breeding has been brought about primarily by improvement in plant type. Further
212 Gurdev S Khush

improvement will be possible by specific selection for tim traits enumerated. The
inheritance of most of these traits is not clearly understood and genes controlling
these traits and other desirable plant characteristics are likely to have both
desirable and undesirable linkages. Any breeding approach that will allow
retention of desirable genetic linkages, break the undesirable (repulsion phase)
linkages and improve the selection efficiency in discriminating between high- and
low-yielding genotypes should be helpful in developing varieties with higher yield
potential. Innovative breeding "methods such as heterosis breeding, recurrent
selection, biparental mating and disruptive mating, and the anther culture method,
aided by modified selection and evaluation procedures, should help meet the
desired objectives.
Assuming that some undesirable genetic linkages limit the yield potential of the
true breeding lines developed through conventional breeding, the F~ hybrids
should help overcome the effect of undesirable linkages by bringing the desirable
dominant genes from two parents together. This approach has been successfully
employed in China (Lin and Yuan 1980) and its utility in the tropics is being
explored (Virmani et al 1981).
Recurrent selection is used primarily to promote recombination and to increase
the frequencies of favorable genes for quantitatively inherited traits. It is cyclic,
encompass!ng the two phases of plant breeding: a) selecting a group of genotypes
that possess favorable genes, and (b) mating the selected genotypes to obtain
genetic recombination (Frey 1982). The applicability of this technique to
self-pollinated species has been limited because of technical problems of
intermating. However, availability of monogenic male sterility (Singh and Ikehashi
1981) in rice variety IR36 has made it possible to use the recurrent selection
technique for developing varieties with higher yield potential.
Biparental Or disruptive mating schemes can also be employed to overcome the
problem of adverse linkages. Biparental mating involves intermating of selected
plants in F2 so as to accumulate favorable genes and to break linkages, thereby
releasing a greater reservoir of genetic variability to enable the breeder to exercise
selection (Joshi 1979). Disruptive mating involves the intermating of unlikes in
segregating generations. It leads to greater opportunity for crossing over, which
releases latent variation by breaking repulsion phase linkages (Thoday 1960).
Plant breeders are often faced with the low efficiency of visual plant selection for
yield in the early segregating generations. Heterogenous soil fertility conditions,
intergenotypic competition due to density of the population, and micro and
macroenvironmental effects are some factors responsi.ble for low selection
efficiency. Dominance and epistatic variance in the segregating populations reduce
selection efficiency. Cultural methods such as high fertilizer rates and wider
spacing, which allow for maximum expression of genotypes for yield and reduce the
w~riability due to environnaental factors, .maY be employed as aids'to selection for
higher yield potential. The haploid method using anther culture is also useful for
improving the selection efficiency for yield and other traits of low heritability. In
the doubled haploid lines there is more additive genetic variance as compared to
the conventional F2 and F3 generations. In addition, dominance variance is
eliminated. In F 3 and F4, additive and dominance effects contribute to phenotypic
differences between individuals, whereas variation in doubled haploid progeny is
only due to microenvironmental effects (Snape 1982). Thus, selection efficiency in
Rice breeding: Past, present and future 213

tile doubled haploid population is likely to be higher when there is greater


dominance variation in the cross.
Improvements in the yield potential of rice to date have been brought abOut
primarily through selection for improved plant type following conventional
methods of pedigree or backcross breeding. However, for further progress in
increasing the yield potential, conventional breeding methods must be sup-
plemented with the innovative breeding techniques discussed earlier.
Grain quality:' Most improved varieties have excellent grain size, shape, and
appearance, and good milling recovery. But most have high amylose con'tent and
low gelatinization temperature of the grain starch. It is now obvious that in the
tropics and subtropics, there is universal preference :for rices with intermediate
amylose and intermediate gelatinization temperature. Fortunately, there seems to
be no genetic barrier to combining these quality traits with high yield or other
adaptability traits. Wide-scale acceptance of IR64 as a high quality rice fortifies our
conviction that rices with intermediate amylose content and intermediate gelati-
nization temperature have superior palatability. Rice breeders should, therefore,
endeavor to incorporate these desirable grain quality features in the future rice
varieties.
Aromatic rices are valued by consumers and command higher prices. Yet this
trait has not been incorporated into any of the improved rice varieties. Basmati
rices of India and Pakistan have grain, elongation traits in addition to aroma, and
are sold at premium prices in national and international markets. Despite
concerted efforts by several rice improvement programs during the last 20 years,
high-yielding varieties with Basmati grain quality have not been developed. The
challenge of developing high-yielding Basmati is a major one for rice breeders. The
methods discussed earlier may prove useful in breaking the undesirable linkages
that are h,lmpering progress in this area.
Yield stability: Yield stabi'lity,depends on multiple resistance to major diseases and
insects. However, resistant varieties do not remain resistant forever. The useful life
of a resistant variety is 5-10 years. We must endeavor to develop varieties with
durable resistance. Polygenic resistance is assumed to be more durable. Recurrent
selection procedures discussed earlier are helpful in accumulating polygenes :for
resistance in the same genotype. Thus, more attention will have to be paid to
developing varieties with more durable resistance.
Wild species of rice are a rich source of genes for disease and insect resistance.
However, this resource has not been exploited for rice improvement. We have
recently succeeded in transferring genes for resistance to brown planthopper and
whitebacked planthopper from Oryza officinalis to O. sativa across crossability and
recombination barriers (Khush and Jena 1987). Other wild species also have useful
genes for resistance and we may have to use the techniques of biotechnology, such
as embry o rescue and protoplast fusion, to move these genes from the wild species
to cultivated rice.
Through genetic engineering techniques, it is now possible to move genes from
unrelated organisms to crops of economic importance. Transfer of the Bt gene from
Bacillus thuringiensis to tobacco is an Qutstanding example of the potential of
genetic engineering techniques in crop improvement. The Bt gene is expressed in
214 Gurdev S Khush

the genetically engineered pla~lts and produces the toxin which kills lepidopteran
insects (Vaeck et al 1987). It is known that some plants such as neem (Azadirachta
indica) produce chemicals that have insecticidal properties. Can we clone the neem
gene or genes responsible for the production of these chemicals and transfer them
to rice through genetic engineering techniques?
To date, sources of resistance to some important diseases such as sheath blight
have not been found, although a large number of germplasm entries have been
screened. Development of varieties resistant to sheath-blight is yet another
challenge to plant breeders.
Shorter growth duration: Several varieties with a growth duration of 110 days have
been developed whose yield potential is comparable to that of varieties with a
130-day growth duration. Can we develop varieties with a 90-95 clay growth duration
without reduction of yield potential? It is obvious that we will have to select geno-
types with very fast growth rates so that they are able to produce the same biomass
in fewer days. It has been observed that short-duration varieties have weaker stems.
Can the linkage between short growth' duration and straw weakness be broken?

Varietal development for unfavorable environments

Unfavorable environments (rainfed lowland, upland deepwater, tidal wetland) are


characterized by variable water regimes, occurrence of drought, submergence,
waterlogging, and soil toxicities or deficiencies. Progress in developing improved
varieties for these environments has been extremely slow because:
The ideal high-yielding plant type for various unfavorable environments is still
not clearly defined.
Adaptability of the genotypes for specific rice culture is most important. Hence
it is essential to retain, as far as pdssible, the adaptability traits of the locally
adapted varieties.
Adaptability and productivity traits are sometimesnegatively correlated, hence
the problem of repulsion phase genetic linkages has to be Overcome.
Inheritance of adaptability traits such as drought-tolerance, submergence
tolerance, elongation ability, and tolerance to mineral stresses is not adequately
understood. Very little information is available oh the mechanisms of tolerance to
these stresses.
* Selectionfor local adaptability can be done only by screening and evaluating the
breeding material in target areas.
Available screening procedures for selecting for tolerance to various stresses do
not give consistent results.
The generation advance is slower because of the complexity of the evaluation
procedures and involvement of photoperiod sensitivity in some populations.
Suitable donors for certain stresses have not been identified.
To develop improved germplasm for the unfavorable environments, the
following strategies will have to be followed.
(i) Definition and characterization of the major favorable rice cultural types and
identification of the representative sites for selection and evaluation of breeding
material. Considerable progress has been' made in these directions but more needs
to be done (Khush 1984).
Rice breeding: Pas:t, present arm future 2115

(ii) Identifichtion of donors for specific traits and varieties, adapted to wtrious
target environments.
(iii) If improvement is desired in a few traits, backcross or mutation methods of
breeding may be employed using a locally adapted variety as the base.
(iv) If a large number of traits are to be improved simultaneously, the conventional
pedigree method of breeding for handling crosses between locally adapted
materials and donors for traits to be incorporated should be adopted.
(v) Population improvement methods of breeding using inale sterile lines and
facilitated recurrent selection should be employed to promote recombination and
increase the frequency of favorable genes in the breeding populations.
(vi) Attempts should be made to improve the harvest index by selecting genotypes
with larger sink size.
(vii) The Rapid Generation Advance (P,GA) technique could be used extensively
for selected crosses involving photoperiod-sensitive parents.
(viii) Tissue culture techniques may be employed to exploit somaclonal variation
to select for stres.s tolerance.
(ix) Wide hybridization may be employed to incorporate traits for stress tolerance
from the wild germplasm.
(x) A shuttle breeding approach may be adopted for screening and evaluating
breeding material in cooperation with scientists working in target areas.

Conclusions

The Green Revolution technology, centered on high-yielding, disease- and


insect-resistant rice varieties, has revolutionized rice production since the late
1960s. Many countries in the rice belt of Asia, which used to import large quantities
of rice, have become self-sufficient and have some surpluses to export. As a result,
rice prices on the international market and in the domestic markets of many
countries have fallen, thus helping the purchasing power of weaker sections of
these societies. The consequent improvement in food security has led to political
stability and allowed the governments of the developing countries to pay more
attention to the pressing needs of economic development.
Population growth is continuing at more than 2% annually in many developing
rice-growing countries. The demand for rice is likely to exceed supply by the year
2000. To feed this growing population, the growth rate of rice production needs to
accelerate further. For this we need varieties with higher yield potential, greater
yield stability, shorter growth duration, and superior grain quality. Innovative
breeding methods and the emerging techniques of biotechnology mt~st supplement
the conventional breeding methods in achieving the future rice breeding goals.
Rice breeding today is. an international effort, involving scientists worldwide.
IRRI is supported by an informal organization of 34 donor agencies called the
Consultative Group on International Agricultural Research (CGIAP.). The Govern-
ment of Japan is the third largest donor to the CGIAR. Numerous Japanese
scientists have made notable contributions to rice science and improvement. Half
of the world's yearly scientific literature on rice science is published in Japan.
The recognition of IRRI's work by the awarding of the 1987 Japan Prize will help
generate additional support for rice improvement in the developing nations. This in
216 Gurdev S Khush

t u r n will h e l p a c h i e v e t h e n o b l e g o a l s o f " P e a c e a n d p r o s p e r i t y f o r a l l " s e t b y t h e


Japan Science and Technology Foundation.

References

Borgstrom G 1972 The hungry planet (New York: MacMillan)


Frey K J 1982 Breeding approaches for increasing crop yields, Paper presented at the International Rice
Research Conference, [RI~,I, Los Bafios
Gonzales L A and Unroll D L 1985 Rice production systems: Issues to be considered. Int. Rice
Commtttt. News'l: 3 4 : 1 9 - 4 8
IFf:'RI 1977 Food needs of developing countries: Projections of production and consumption to 1990,
International Food Policy Research Institute, Washington, DC
[Rill 1982 IR36, the world's most popular rice, International Rice Research Institute, Manila,
Philippines
Joshi A B 1979 Breeding methodology for autogamous crops, b~dian J. Genet. Plant Breed. 39:567-578
K,hush G S 1977 Disease and insect resistance in rice. Adv. Agtvtt. 29:265-341
Khush G S 1984 Ternfinology for rice growing environments. In Telvninology ]or rice growing
environtllelltS (Manila: IRRI) pp. 5-10
Khush G S 1987 Development of rice wuieties suitable for double cropping. In lnternationalsymposium
on technology for double cropping o,['rice in tropics (Yatabe, Tsukuba, Japan: Tropical Agriculture
Research Center) pp. 235-246
Khush G S and Jena K K 1987 Transfer of genes from O. oJ]}'cinulis to cultiwited rice across crossability
and recombination barriers. Agron. Abstr. for 1987 (In press)
Lin S C and Yuan L P 1980 Hybrid rice breeding in China. In hmovative approaches to rice breeding.
(M~mila: IRRt) pp. 35-51
Paddock W and Paddock P 1967 Famine-1975! (Boston-Toronto: Little Brown and Company)
Singh R 3 and Ikehashi t-I 1981 Mon0genic male sterility in rice: induction, identification and
inheritance. Crop Sci. 21:286-289
Snape J W 1982 The use of doubled haploids in plant breeding. In hlduced variability in plant breeding.
International Symposium on Mutatiot~s and Polyploidy (Eucarpia, Wageningen, The Netherlands) 19:
143
Thoday J M 1960 Effects of disruptive selection. III. Coupling and repulsion. Heredity 14: 35-4~
Vaeck M, Reynaerts A, Hofte H, Jansens S, de Beuckeleer M, Dean C, Zabeau M, Van Montagu M
and Leemans J 1987 Transgenic plants protected from insect attack. Nat'tre (London) 328:33-37
Virmani S S, Chaudh~lry R C and Khush O S 1981 Current outlook on hyblid rice. Oryza 18:67-84

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