Journal of Integrative Plant Biology 2007, 49 (6): 805−810

.Invited Review.

Super Hybrid Rice Breeding in China:

Achievements and Prospects
Shi-Hua Cheng*, Li-Yong Cao, Jie-Yun Zhuang, Shen-Guang Chen, Xiao-Deng Zhan,
Ye-Yang Fan, De-Feng Zhu and Shao-Kai Min

(State Key Laboratory of Rice Biology/National Center for Rice Improvement, China National Rice Research Institute,
Hangzhou 310006, China)

Abstract

Hybrid rice has contributed greatly to the self-sufficiency of food supply in China. To meet the future demand for
rice production, a national program on super rice breeding was established in China in 1996. The corresponding
targets, breeding strategies and most significant advances are reviewed in this paper. New plant type models have
been modified to adjust to various rice growing regions. In recognition of the importance of applying parents with
intermediate subspecies differentiation in increasing F 1 yield, medium type parental lines were selected from
populations derived from inter-subspecies crosses with the assistance of DNA markers for subspecies
differentiation. Results also indicate that a substantial increase of biomass is the basis for further enhancement of
the grain yield potential, and amelioration of leaf characteristics is helpful in increasing the photosynthetic rate.
Thirty-four super hybrid rice varieties have been released commercially, growing in a total area of 13.5 million hm2
and producing 6.7 thousand million kg more rice in 1998–2005. Although remarkable progress has been made in
super hybrid rice breeding in China, selections on the root system and integration of more biotechnological tools
remain a great challenge.

Key words: biomass; DNA marker; harmonious plant type; medium type parents; super hybrid rice.

Cheng SH, Cao LY, Zhuang JY, Chen SG, Zhan XD, Fan YY, Zhu DF, Min SK (2007). Super hybrid rice breeding in China: Achievements
and prospects. J. Integr. Plant Biol. 49(6), 805−810.
Available online at [Link]/links/toc/jipb, [Link]

China’s Future Demand for High Yield Rice more than 60% of the population and contributing nearly 40%
of total calorie intake. China has acted as a pioneer in rice
Rice (Oryza sativa L.) is a staple food crop in China, feeding breeding in most of the last five decades. With the wide adop-
tion of semidwarf varieties since the end of the 1950s, rice
yield in China increased from 2.0×103 kg per hm2 in the 1960s
Received 20 Dec. 2006 Accepted 10 Mar. 2007 to 3.5×103 kg per hm2 in the 1970s. Subsequently, hybrid rice
Supported by the National Program on Super Rice Breeding, the Ministry of that has a yield advantage of 10%–20% over the conventional
Agriculture, National High-Tech and Development Program of China (863), varieties was developed in 1973 and released commercially in
the Ministry of Science and Technology and Provincial Program (8812), 1976. Together with the application of modern cultivation
Zhejiang Province, China. technology, the rice yield has risen to about 6.0×10 3 kg per
Publication of this paper is supported by the National Natural Science hm2.
Foundation of China (30624808). The annual acreage under hybrid rice has increased from
*Author for correspondence. 0.14 million hm2 in 1976 to about 15 million hm2 in recent years,
Tel: +86 (0)571 6337 0192; accounting for more than 50% of the total rice planting areas in
Fax: +86 (0)571 6337 0989; China. It was estimated that hybrid rice in China was grown in a
E-mail: <shcheng@[Link]>. total area of 370 million hm2 from 1976 to 2005 and increased the
© 2007 Institute of Botany, the Chinese Academy of Sciences grain production by nearly 450 thousand million kg (Cheng et al.
doi: 10.1111/j.1672-9072.2007.00514.x 2006). This contributed a great proportion to the self-sufficiency
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806 Journal of Integrative Plant Biology Vol. 49 No. 6 2007

of the food supply in China. However, the yield ceiling wit- addition to desirable grain quality and pest resistance, the yield
nessed in various crop species (Mann 1999) has also been targets of super hybrid rice under favorable conditions are
encountered in the rice production in China since 1990 (Cheng expected to meet the future requirement (Table 2).
et al. 2004a, Yang et al. 2006). In addition, arable land available According to the ecological and cropping conditions of dif-
for rice cultivation has decreased during the last three decades, ferent rice growing regions in China, a set of quantified models
reducing rice planting area from 36.2 million hm 2 in 1976 to of the plant type for super rice has been formulated. For
26.5 million in 2003. As a consequence, rice production in China instance, the plant type model of indica hybrid rice in the Yangzte
fell from a high record of 200.7 thousand million kg in 1997 to River Valley rice regions is formulated as follows.
160.7 thousand million kg in 2003. · Medium tillering capacity with 12−15 panicles per plant
To ensure food security for the increasing population, raising · 115−125 cm plant height
the yield ceiling of rice remains a priority in China. Based on the · Droopy panicle with erect, longer and slight rolling flag leaf
per capita rice consumption of 150 kg and a stable rice crop- · 190−220 grains per panicle
ping area of 31.57 million hm2, the rice yield should be increased · Vigorous root system
to 6.885×103 and 7.845×103 kg per hm 2 in 2010 and 2030, · Multiple resistance to diseases and insects
respectively (Table 1) (Cheng et al. 2005). However, an in- · 135−140 d growth duration
crease in rice yield has become more and more difficult. · Harvest index of 0.55
· 12−15×103 kg/hm2 yield potential

Targets of the Super Rice Breeding Program

in China Breeding Strategies for Super Hybrid Rice

Based on the experiences of high-yield rice breeding in China

Increasing the genetic diversity of parents
and new plant type rice (super rice) breeding at the Interna-
tional Rice Research Institute (IRRI), a special collaborative re- Importance of applying parents with intermediate
search program on super rice breeding was established by the subspecies differentiation in increasing F 1 yield
Ministry of Agriculture, China in 1996. The program mainly fo- The majority of inbreeds and hybrids of rice grown in China
cuses on the breeding of super hybrid rice, which is defined have few ancestors according to the pedigree analysis. As
as the varietal type that combines the harmonious plant type indicated by a number of DNA marker-based analyses (Yang et
with heterosis and achieves super high yield through hybrid- al. 1994, Zhuang et al. 1998), narrowing genetic variability
ization between indica and japonica (Cheng et al. 1998). In among modern semi-dwarf rice varieties and their primary par-
ents are strongly evident. The insufficient genetic diversity is
recognized as a major cause for the yield ceiling in hybrid rice
Table 1. Future demand for rice production and yield in China
that has appeared for nearly 20 years.
Total output Increase Yield Increase
Year Rice is generally classified into two subspecies, indica and
(×109 kg)a (%)b (kg/hm2) (%)b
japonica. Inter-subspecies heterosis has long been attempted,
2010 217.5 19.18 6 885 16.3
but it is difficult due to unharmonious genetic backgrounds be-
2030 247.5 35.62 7 845 32.6
a tween two subspecies. To test the relationship between sub-
Estimated based on the per capita rice consumption of 150 kg and
species differentiation of the parental lines and the F 1
the rice cropping area of 31.57 × 10 6 hm 2 ; bIn comparison with
productivity, a double haploid (DH) population was developed
1995.
from an inter-subspecies cross Teqing (indica)/02428
(japonica). DH plants with different subspecies differentiation
Table 2. Yield target of different types of super hybrid rice varieties were used as restorer lines to testcross indica CMS
(Ministry of Agriculture, China, 1996) (cytoplasmic male sterility) maintainer line Xieqingzao B and
Yield target (×10 3 kg/hm 2) a Increase (%)b indicalinous CMS maintainer line 064 B, respectively. The cor-
Year
EI (Y) SI & J LI relation between subspecies differentiation indexes on DH
2000 9.75 10.50 9.75 15 plants and the yield in F1 plants was analyzed. Results indicate
2005 11.25 12.00 11.25 30 that hybrids from the indicalinous or japonicalinous DH plants
a
Performance at two sites, 6.67 hm2 or more at each site, successive (differentiation index 11–15) have yield advantages over other
two years. bYield advantage over the control variety in multi-loca- F1 plants (Figure 1). Therefore, selection of medium type par-
tion trials. EI (Y), early season indica rice in middle and lower ents from populations derived from inter-subspecies crosses
reaches of Yangtze River. SI & J, single season indica or japonica has attracted great attention in the super hybrid rice breeding
rice. LI, late season indica rice. program in China.
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Super Hybrid Rice Breeding in China 807

Selection of DNA markers for subspecies polymorphism) markers were first employed (Qian et al. 1995),
differentiation in rice but technical complexity and cost made it unfeasible for use in
As mentioned above, subspecies classification of rice culti- the general breeding programs. SSR (simple sequence repeat)
vars and selection of parental lines with intermediate subspe- markers, which are technically easy and less expensive to
cies differentiation is a subject of importance for rice breeders. use, were screened when rice microsatellite framework map-
Cheng’s subspecies differentiation index (Cheng 1985) based ping became available (Chen et al. 1997). A set of 21 SSR
on six morphological and biochemical traits was extensively markers distributing on the 12 rice chromosomes were se-
used previously. Since DNA marker offers a new approach lected (Fan et al. 1999) and adopted by the breeders for paren-
that is more effective and reliable for the subspecies classifi- tal selection.
cation in rice (Wang and Tanksley 1989; Neeraja et al. 2003),
efforts have been made to screen DNA markers showing dif- Development of medium type restore lines in
ferent patterns between subspecies. Two sets of DNA mark- subspecies differentiation
ers for subspecies differentiation in rice were identified (Table Since last decade, the development of medium type restorer
3) (Cheng at al. 2004b). RFLP (restriction fragment length lines in subspecies differentiation has become more and more
applied in the hybrid rice breeding programs in China. In the
indica rice growing regions of China the breeders adopted the
methodology of introgressing japonica blood into an indica rice
background to develop indicalinous germplasms, and in the
japonica rice growing regions introgressing indica blood into
japonica rice background to develop japonicalinous
germplasms. So far, a set of indicalinous or japonicalinous
germplasms has been deliberately developed, of which some
have been successfully used in the breeding of super hybrid
rice. For instance, R9308, an indica restorer line from an inter-
subspecies cross, was selected and successfully used in the
development of super hybrid rice Xieyou 9308 by CNRRI (Figure
2). It is estimated that there are 25% japonica genetic compo-
nents in R9308. Its hybrid Xieyou 9308 is estimated to have
12.5% japonica genetic components and has a good combina-
tion of super high yielding capacity with harmonious plant type
Figure 1. Relationship between subspecies differentiation index of (Figure 3). This hybrid was released commercially in Zhejiang
the paternal parents and grain yield per plant of their F 1 hybrids. Province in 1999 (Cheng et al. 2005) and has a total planting
area of over one million ha. Another example is the two-line
super indica hybrid Liangyou Pei 9 developed by Jiangsu Acad-
Table 3. Chromosomal distribution of DNA markers for subspecies
classification of rice varieties
Chrom-
RFLP (28 markers) SSR (21 markers)
osome
1 RG101 RG345 RG462 RG472 RM23 RM259
2 RG171 RG256 RG322 RM29 RM250
3 RG482 RG96 RM16 RM251
4 RG214 RG620 RM226
5 RG207 RG474 RM13
6 RG64 RZ828 RM217
7 RG351 RG511 RM18 RM234 RM248
8 RG978 RZ562 RM25
9 RG553 RG570 RG667 RM245 RM205
10 RG752 RZ811 RM258 RM228
11 RG167 RM202
12 RG81 RG543 RG958 RM4 RM20 RM247
RFLP, restriction fragment length polymorphism; SSR, simple se- Figure 2. Development of medium type restorer line R9308 and its
quence repeats. super hybrid Xieyou 9308.
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808 Journal of Integrative Plant Biology Vol. 49 No. 6 2007

at Xinchang County, Zhejiang Province, the grain yield of super

hybrid rice Xieyou 9308 was 12.23×103 kg/hm2, being 38.2%
higher than that of the control hybrid Xieyou 63. Furthermore,
the biomass of Xieyou 9308 was 42.7% higher than that of
Xieyou 63, but the harvest index was 3.9% lower than that of
Xieyou 63, indicating that the increased grain yield of Xieyou
9308 is associated with the increased biomass (Zhai et al. 2002).
It can be concluded that the key to further increasing the
grain yield potential is to increase biological yield while main-
taining the current harvest index. Moderate increase of plant
height had been considered a very promising way of increas-
ing biological yield, but lodging was often associated with the
increase of plant height. Recently, increasing biological yield
through developing the short stature with stronger tillering
capacity, thicker stems and thicker and more erect leaves were
Figure 3. Field performance of Xieyou 9308 with harmonious plant proposed as an alternative approach.
type, in Xinchang, Zhejiang Province, China.
Amelioration of the leaf stature and quality to increase
photosynthetic rate
emy of Agricultural Sciences in collaboration with Hunan Hy- Currently, some rice varieties in high-yielding cultivation have
brid Rice Research Institute. Its female parent Peiai64S is a leaf area indexes of 8–10 and this seems to be the maximum. In
TGMS (thermo-sensitive genetic male sterility) line with indica, order to increase the number of spikelets and the number of
temperate japonica and tropic japonica ancestries. filled grains per panicle, it is necessary to ameliorate the leaf
stature and quality. It has been well demonstrated that erect
Construction of harmonious plant type and slight rolling uppermost three leaves favor the full use of
light energy after heading, by promoting CO2 diffusion, increas-
Role of substantial biomass production in increasing ing photosynthetic rate on the back of leaves, accelerating the
grain yield in rice increase in biological yield, mitigating the conflict between
It has been known that yield increase of rice through replace- panicle number and size, and improving lodging tolerance of
ment of tall inbred cultivars with semidwarf varieties is due to the rice plant. For instance, the photosynthetic rate of a flag
the increase of harvest index, while the yield advantage of leaf in the super hybrid Xieyou 9308 and control hybrid Shanyou
hybrid rice is attributed to the increase of biological yield (Table 63 are similar in the up-face, but it is 21.6% higher in Xieyou
4). It has also been recognized that enhancement of harvest 9308 than in Xieyou 63 in the down-face, which is attributed to
index is of great difficulty. In an experiment under the special higher leaf rolling in Xieyou 9308 (Table 5) (Zhu et al. 2001).
ecological conditions at Yongsheng County, Yunnan Province,
China, the grain yield, biological yield and harvest index of in-
dica hybrid Shanyou 63 was 15.27×103 kg/hm2, 28.29×103 kg/ Outcome of Super Hybrid Rice Breeding in
hm2 and 0.54×103 kg/hm2, respectively. Compared with the per- China
formance of Shanyou 63 under the usual ecological conditions,
the grain yield and biological yield were much higher, whereas Following the breeding strategies and plant models designed
the harvest index was almost identical (Yang et al. 2006). Simi-
lar results were obtained when the comparison was made Table 5. Comparison of leaf rolling and photosynthetic rate in super
between the super rice hybrid and the control hybrid in usual hybrid Xieyou 9308 and control hybrid Shanyou 63
conditions. In an experiment under usual ecological conditions Xieyou 9308 Shanyou 63
(Super hybrid) (CK)
Table 4. Biomass base of increasing grain yield in different type of Leaf rolling (%)
varieties Flag leaf 15.6 9.7
Grain yield Biomass Harvest Second leaf 11.0 6.2
Year Varietal type
(×103 kg/hm2) (×103 kg/hm2) index Third leaf 10.3 4.5
1940s–60s Tall inbred 4.23 11.00 0.385 Photosynthetic rate in flag leaf (μmol CO 2.m−2.s−1)
1960s–80s Semi-dwarf inbred 6.43 11.81 0.545 Up face of the leaf 22.7 23.2
1970s–90s Hybrid 8.20 15.04 0.545 Down face of the leaf 22.5 18.5
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Super Hybrid Rice Breeding in China 809

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