Rice Breeding: Past, Present and Future: (IRRI)
Rice Breeding: Past, Present and Future: (IRRI)
Rice Breeding: Past, Present and Future: (IRRI)
GURDEV S KHUSH*
Department of Plant Breeding, International Rice Research Institute, P.O. Box 933,
Manila, Philippines
*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.
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.
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)
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.
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
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
Reaction"
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 -
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
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.
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
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
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.
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
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
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~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
I00
83
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2
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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
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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
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210 Gurdev S Khlxsh
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1960 '65 '70 '75 '80 '8! Figure 10. Rice export price at
Yeor Bnngkol~ (deflated)
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
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
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?
(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
References