Indian J. Genet.

, 75(3): 279-293 (2015)

DOI: 10.5958/0975-6906.2015.00045.0

Review article

From concept to field: evolution of hybrid pigeonpea technology in

India
1
K. B. Saxena *

International Crops Research Institute for the Semi-Arid Tropics, Patancheru 502 324, Telangana

(Received: February 2015; Revised: June 2015; Accepted: July 2015)

Abstract in maize have saved the world from malnutrition and

The role of hybrid vigour in the enhancement of productivity
hunger. The maize yields in the USA increased five
is well recognized. Its application, however, has been folds from a meager 25 bushels/acre in 1930 to 140
restricted to a few crops and legume breeders could never bushels/acre in1998 (Trayer, 1991; Crow, 1998). This,
take its advantage due to various seed production issues. beyond doubt, happened due to untiring and continuous
Recently, plant breeders have succeeded in creating the efforts of scientists in developing improved hybrids
world’s first commercial hybrid in a food legume, popularly
and their production technology. The process of
known as red gram or pigeonpea [Cajanus cajan (L.) Millsp.].
This was possible due to success in breeding a stable recombination and selection that contribute to yield
cytoplasmic nuclear male sterility (CMS) system and and stability is still continuing and contributing towards
exploitation of its partial natural out-crossing. This further enhancement of yield and adaptation of maize
development has provided golden opportunities to break hybrids in different parts of the world. The phenomenal
the decades-old yield plateau in this crop. Among the
success of hybrid maize led breeders to use this
hybrid combinations evaluated, ICPH 2671 and GTH 1 were
found most promising. In the on-farm trials ICPH 2671 technology in other crops like cotton, sorghum, pearl
recorded 46.5% superiority over the best available cultivar millet, sunflower, brassica, safflower and various
Maruti; while GTH 1 was 33.9% superior to the control, and horticultural crops. At present efforts are also being
it was released in Gujarat state in 2004; but for some reasons made to bring together the genes responsible for
associated with its stability of fertility restoration it failed
dwarfing and hybrid vigour in wheat and rice with a
to reach the level of commercialization. The outstanding
performance of hybrid ICPH 2671 led to its release by both, target of harvesting more grains per unit of land area.
a private seed company as ‘Pushkal’ and by the state After the adoption of hybrid technology in various crop
Government of Madhya Pradesh as ‘RV ICPH 2671’ in 2010. species, now it is the turn of grain legumes. This
The journey of the evolution of hybrid pigeonpea endeavour started with research in faba bean in the
technology was fascinating and challenging. It took a long west (Bond et al. 1966) and pigeonpea in India
time of over 39-years from its conceptualization to finally
reaching Indian farmers. The impact of this breakthrough
(Saxena, 2008). Efforts are also being made to develop
was visible when two more hybrids ICPH 2740 and ICPH hybrid breeding technology in soybean (Palmer et al.
3762 were recently released in India. Since it is a path 2010). The commercial success in this direction,
breaking research among food legumes, an attempt has however, has come only from pigeonpea or red gram
been made here to archive the milestones of this historic [Cajanus cajan (L.) Millsp.], a popular protein-rich pulse
plant breeding event.
crop of Asia, Southern and Eastern Africa, and the
Key words: Cajanus cajan, cytoplasmic nuclear male Caribbean islands. The recent release of three
sterility, hybrid, out-crossing, pigeonpea commercial hybrids in India has established the hybrid
technology in pigeonpea also. This paper briefly reviews
Introduction the important research and developmental initiatives
Commercial exploitation of two genetic phenomena - undertaken by International Crops Research Institute
the dwarfing genes in rice and wheat and hybrid vigour for the Semi-Arid Tropics (ICRISAT) and Indian Council

*Corresponding author’s e-mail: [email protected]

1
Present address: Villa 17, NMC Housing Complex, Al Ain, Abu Dhabi, U.A.E.
Published by the Indian Society of Genetics & Plant Breeding, F2, First Floor, NASC Complex, PB#11312, IARI, New Delhi 110 012
Online management by indianjournals.com
280 K. B. Saxena [Vol. 75, No. 3

of Agricultural Research (ICAR) in the last 39 years natural out-crossing of the crop for population
that led to the successful development and adoption improvement; and in spite of about 2% yield gain per
of hybrid pigeonpea technology. cycle of selection in Kenya (Onim, 1981), this approach
did not take-off as a standard breeding method in
Background information pigeonpea. Consequently, pure line breeding remained
Pigeonpea is a drought tolerant short-lived perennial a major instrument for the genetic improvement of this
pulse, cultivated on over 4 m ha (IIPR, 2013) at pulse and in the past 4-5 decades over 100 pure line
subsistence level in India, mainly as an intercrop. Its cultivars were released (Singh et al. 2005).These
decorticated dry splits, whole grains, and fresh peas cultivars although exhibited significant genetic
are consumed with cereals as a protein supplement. advances for traits such as earliness, plant type,
Besides fixing atmospheric nitrogen, it is also disease resistance and market-preferred traits such
recognized for its ability to release soil-bound as seed size and colour; but without any marked
phosphorus, adding organic matter, and recycling of improvement in the productivity that remained
valuable nutrients in the soil (Saxena, 2008). unchanged over the past half century at around 700-
800 kg/ha (Fig. 1).
Over a half-century of productivity stagnation in
pigeonpea Yield stagnation in pigeonpea has been a matter
of concern, particularly in view of increasing population
In India the formal genetic improvement of pigeonpea and decreasing per capita protein availability in the
started in 1931 with pure line selection within landraces country, which is about 25% short of the prescribed
for disease resistance, plant type and seed yield standard of 42 g protein/head/day (NIN, 2010).
(Ramanujam and Singh; 1981). Green et al. (1979, According to Shalendra et al. (2013) the pulse
1981), while reviewing the accomplishments and production has registered an impressive growth of
constraints of the global pigeonpea improvement 3.47% during 1990-2009; but it was insufficient to take
programmes, concluded that almost all the breeding care of the national requirements. Further, high
methods traditionally recommended for self-pollinated population growth and low crop productivity were
crops have been tried for the genetic enhancement of considered the two key constraints in meeting the
pigeonpea with limited success. Khan (1973), Sharma challenges of the national nutritional security. In this
and Green (1977) and Onim (1981), however, had context it is reasonable to assume that a quantum
other ideas and decided to use the inherent partial jump in productivity is the only viable solution and

Fig. 1. Area, production and yield (kg/ha) of pigeonpea in India

August, 2015] Hybrid pigeonpea review 281

hybrid technology offers great promise. The recent and low temperature has been found to offset the effect
releases of three commercial hybrids have generated of photo-period to some extent (Turnbull et al. 1981).
a lot of optimism among pigeonpea scientists and policy An important aspect of photo-period sensitivity is
makers towards breaking the decades-old productivity induction of early flowering in the sensitive materials,
barrier in this crop. when sown under inductive (short day) conditions. This
not only has telescopic effect on flowering time but
Significance of maturity groups in pigeonpea also reduces plant height and biomass (Spence and
Perhaps pigeonpea is the only pulse crop which is Williams 1972). This helps in the cultivation of
credited to have the largest genetic diversity for pigeonpea as a rabi (post-rainy) season crop for
maturity among cultivated types with almost a generation advance and seed multiplication of elite
continuous variation from 75 to >250 days. Broadly, it lines.
has five maturity groups – super early (<90 days), Natural out-crossing
extra early (110-120 days), early (140-160 days),
medium (180-200 days), and late (> 250 days). Of Most pulses are known for their cleistogamous flowers
these, the super early group is of recent origin (Vales and self-pollination. It is, however, not entirely true for
et al. 2012) while the rest being there for a long time. pigeonpea; and the first report on its partial natural
Each maturity group has its own significance in relation cross-pollination was published by Howard et al. (1919).
to cropping system. This classification is so strong Subsequently, a number of reports have appeared in
that one group cannot easily replace the other, as far literature (see review by Saxena et al. 1990). Some
as its adaptation is concerned. The early maturing efforts have also been made to understand the factors
pigeonpea has a number of production niches (Saxena responsible for out-crossing in this crop; and frequent
et al. 2014a), while the medium and long duration types insect visits were recognized as the prime cause
are adapted to subsistence agriculture in intercropping (Pathak 1970; Williams 1977; Onim 1981; Zeng-Hong
systems. The hybrid breeding programmes, therefore, et al. 2011). In these studies over two dozen insect
should be targeted towards specific cropping system species were found foraging on pigeonpea flowers,
and maturity group. To reduce genotype x environment but out-crossing was affected by only a few of them.
interaction and enhance selection efficiency Byth et Williams (1977) reported that in Patancheru (India)
al. (1981) recommended that pigeonpea breeding Megachile bicolor and M. conjuneta were the major
activities should be carried out under the cropping pollinators; while Onim (1981) reported that cross-
system for which the end product (cultivar) is targeted. pollination in pigeonpea was primarily a function of
foraging by Apis mellifera and Megachile species in
Photo-period sensitivity and ratooning Kenya. Recently, Zeng-Hong et al. (2011) reported
Pigeonpea is known to be a quantitative short day that in Yuanmou (China) the insects belonging to
plant. Its perennial growth habit, deep (3-4 m) and Megachile spp., Xylocopa and Apinea spp. were most
strong tap root system help in overcoming short spells frequent visitors to pigeonpea fields (Table 1) and they
of abiotic stresses by regenerating the plant growth, were very active in the collection and transportation
popularly called as ‘ratoon growth’, by utilizing its food of pollen grains from one plant to another, and thereby,
reserves. By nature, pigeonpea plants of all the
maturity groups are capable of ratooning, but the long Table 1. Proportions of four important insect species
duration types ratoon better than early types. The recorded in pigeonpea fields and their activity
on flowers in Yuanmou (China)
perennial growth habit of pigeonpea is also helpful in
hybridizing the diverse (early with late) maturity types. Pollinator Proportion Flowers Stay on
This is possible due to emergence of second and third insect species of insects visited/10 each flower
flushes of flowers within the same crop season present (%) min (sec)
(Saxena et al. 1976). In pigeonpea so far there is no
Megachile spp. 49.04 4.5 6.6
report of the existence of any true photo-insensitive
Xylocopa spp. 11.06 8.9 2.4
genotype. Wallis et al. (1981), however, reported that
the earliest flowering types showed least sensitivity Apinae spp. 10.10 2.5 14.8
to photo-period. Like other crops in pigeonpea also, Catopsilia spp. 7.21 4.0 12.4
the influence of day/night temperature in the Others (15 spp.) 22.59 - -
manifestation of photo-period effects is quite strong Source: Zheng-Hong et al. (2011)
282 K. B. Saxena [Vol. 75, No. 3

affecting cross-fertilization. They observed that the phase deals with the process of achieving the
pollinating insects were more frequent on male fertile breakthrough by breeding two viable CMS systems.
plants with a mean of 4.8 visits/10 minutes as The information about breeding and release of CMS-
compared to 2.8 visits/10 minutes on the male sterile based hybrids and their seed production technology
plants. They attributed this behaviour of the insects has been incorporated in Phase III. In the fourth Phase,
to the presence of (i) chemicals such as flavone and the research and development plans for the production
flavonol, (ii) nectar and (iii) specific scent that is of second generation hybrids with potentially greater
produced by mature pollen grains in the fertile plants. yields have been discussed. The important events of
They further reported that even with 50% less insect each phase along with some pertinent research data
visits, the male sterile plants produced cross-pollinated are illustrated in the following text.
seed yield (384 g/plant) similar to that of more
frequently visited fertile plants (357 g/plant). These Phase I (1974-1990): a giant step towards improving
observations indicated that very high insect activity pigeonpea productivity
may not be essential to produce reasonably good At ICRISAT a broad-based pigeonpea improvement
quantities of hybrid seed. Since there is no wind programme was launched in 1974. The breeding team,
pollination in pigeonpea (Kumar and Saxena 2010), jointly led by Dr. D. Sharma and Late Dr. John M.
the entire cross-pollination can be attributed to insect Green, decided that besides using traditional methods
activity. In this situation it is logical to assume that for breeding high yielding pure line cultivars to meet
the population of pollinating insects cannot be the global needs, some non-conventional breeding
same across the locations and time of the year. approaches should be tried to overcome the constraint
Therefore, a large variation (0-70%) in natural out- of persistent yield plateau. In this context, a decision
crossing has been recorded (Saxena et al. 1990). was made to convert the constraint of partial natural
Besides this, some external factors are also known to out-crossing into an opportunity through heterosis
affect the extent of natural out-crossing. These include breeding. Since this technology required a mechanism
speed and direction of wind (Bhatia et al. 1981), for mass hybrid seed production, as a first step, a
extended periods of stigma receptivity (Dalvi and programme was launched to look for a stable CMS
Saxena 2009), floral morphology of genotype (Byth et source in germplasm that was sown at ICRISAT farm
al. 1982), and immediate surroundings of pigeonpea for a routine assessment and characterization exercise.
plots (R. V. Kumar, personal communication).
The first genetic male sterility (GMS) discovered
Evolution of hybrid pigeonpea technology
A thorough search of over 7000 germplasm accessions
th
Hybrid technology flourished in the 20 century through failed to meet the expectations of finding a cytoplasmic
open-pollinated crops such as maize, pearl millet etc. nuclear male sterility (CMS) for commencing a hybrid
and benefitted millions of farmers; but for pulse breeders breeding programme, but instead, a source of genetic
it remained a challenge. At ICRISAT and ICAR, the male sterility (GMS) was identified in a field collection
hybrid pigeonpea breeding programme started from a from Andhra Pradesh. This GMS line was of medium
scratch and it passed through many cycles of (about 180 days) maturity, susceptible to both fusarium
successes and failures. Finally, the success in this wilt and sterility mosaic diseases. The anthers were
endeavour was achieved, when pigeonpea hybrids GTH translucent and the male sterility was controlled by
1 (in Gujarat) and ICPH 2671 (in Madhya Pradesh) single recessive gene ‘ms1’ and most likely it evolved
were released for cultivation. through spontaneous mutation (Reddy et al. 1978);
and the recessive allele was maintained in the
In this article some major events of research
population through natural out-crossing. After the
and developmental activities that led to the
search of this GMS source, a few more GMS sources
development of hybrid technology are highlighted. The
were identified at ICRISAT and ICAR centres at Akola
author, who was associated with this programme from
and Pantnagar (see review by Saxena et al. 2010) but
the take-off point, has divided this review into four
none of them was used in breeding hybrids.
broad phases for the sake of clarity and has covered
almost entire work of four decades. The Phase I Release of the first GMS-based hybrid ICPH 8
summarize the initial research activities and covers
the period up to the development/release of the first The breeding team knew that GMS was not a perfect
GMS-based pigeonpea hybrid ICPH 8. The second platform for breeding commercial hybrids; but decided
August, 2015] Hybrid pigeonpea review 283

to continue research in order to find answers to basic yielded the control T 21 by a margin of 47.4%. In the
questions such as quantum of hybrid vigour available All India Coordinated Trials also this hybrid exhibited
in the crop and secondly, if the limited natural out- 14% superiority over the best national check UPAS
crossing would be enough to produce large quantities 120. In 1994, another pigeonpea hybrid CoH 1 was
of hybrid seed economically. Therefore, a programme released by Tamil Nadu Agricultural University,
was launched to transfer the male sterility gene to Coimbatore. In 17 on-farm trials CoH 1 recorded 21%
diverse elite lines. higher yield over control VBN-1 (Murugarajendran et
al. 1995). Subsequently, two more pigeonpea hybrids,
By 1979, the first converted early maturing GMS AKPH 4104 (in 1997) and AKPH 2022 (in 1998) were
line ‘MS Prabhat (DT)’ was bred using off-season released for central India by PKV, Akola. These
facility and made ready for use in making the first set hybrids respectively recorded 30% and 64% superiority
of experimental hybrids. At this time the male sterile over the control cultivar (Wanjari et al. 1999).
line was shared with various ICAR centres, Agricultural Interestingly, all the hybrids (non-determinate types)
Universities, and Maharashtra Hybrid Seed Company demonstrated large yield advantages in different zones
(Mahyco) to develop locally adapted hybrid technology. (Table 2), but none of them could reach the level of
The entire hybrid breeding programme was coordinated commercialization, primarily due to the big issue of
jointly by ICAR and ICRISAT. Among the experimental rouging fertile segregants within the female rows.
hybrids tested, a short duration hybrid ICPH 8,
synthesized by crossing MS Prabhat (DT) and an Table 2. List of GMS- based hybrids released in India
advanced breeding line ICPL 161, was found to be the
most promising in station trials conducted from 1981 Hybrid Origin Year Adaptation Maturity Stan-
to 1983. Simultaneously, an exercise began to use zone group dard
hete-
this hybrid combination for developing its large-scale
rosis(%)
seed production technology.
ICPH 8 ICRISAT 1991 Central Early 35
In 1984, ICPH 8 was evaluated along with national
PPH 4 Ludhiana 1994 North west Early 14
check UPAS 120 in All India Co-ordinated Trials
CoH 1 Coimbatore 1994 South Early 21
organized by ICAR. Seed yield of this hybrid over 94
trials, conducted over four years in three climatic CoH 2 Coimbatore 1997 South Early 35
zones, was 34.55% more than the control UPAS 120 AKPH 4104 Akola 1997 Central Early 64
(Saxena et al. 1992). This demonstrated both greater AKPH 2022 Akola 1998 Central Medium 30
yield and stability of ICPH 8 over the inbred cultivars.
With the out-standing performance in the Co-ordinated
To assess the impact of GMS-based hybrid
Trials, the hybrid was promoted by ICAR to pre-release
technology, ICRISAT appointed an ‘impact
on-farm testing in central zone. In this endeavour, the
assessment group’ to record the views of seed
hybrid demonstrated >25% yield advantage in different
companies, seed producing farmers, and hybrid
districts of Maharashtra, Andhra Pradesh, and Gujarat
cultivators in Maharashtra and Andhra Pradesh. The
states. These performances led to the release of ICPH
group concluded that “though the hybrid advantage in
8 by ICAR in 1991 (Saxena et al. 1992). This
pigeonpea is salable and its cost of seed production
endeavour clearly demonstrated the presence of
is also within affordable limits, but the technology itself
commercially exploitable hybrid vigour in pigeonpea.
suffers from a major bottleneck, when it comes to large
In the seed production exercise also, it was observed
scale seed production” Niranjan et al. (1998). The hybrid
that good hybrid yields of the order of 800-1000 kg/ha
advantage recorded in the GMS-based hybrids was
can be obtained through partial natural out-crossing.
encouraging and only seed production issue needed
Release of other GMS-based hybrids by ICAR to be addressed. This was only possible if a stable
cytoplasmic nuclear male sterility (CGMS or CMS),
The primary GMS source (ms 1 ) was also
its maintainers and restorers were discovered.
simultaneously used in other breeding programmes of
ICAR. This exercise resulted in the release of five Phase II (1991-2003): the major breakthrough in
more GMS-based hybrids. In 1993, Punjab Agricultural breeding CMS systems
University, Ludhiana, developed pigeonpea hybrid PPH
In plants the CMS system is primarily conditioned by
4 (Verma and Sidhu 1995). In the state level multi-
cytoplasm containing aberrant mitochondrial DNA,
location trials conducted for two years, PPH 4 out-
284 K. B. Saxena [Vol. 75, No. 3

commonly designated as ‘sterile’ (S) cytoplasm. In population of Cajanus scarabaeoides (L.) Thou., a wild
this case the extra nuclear genome of a genotype relative of pigeonpea. This plant was partially male
interacts with its nuclear genome; and in the presence sterile and in its F2 generation a number of segregants
of recessive non-restoring nuclear alleles (frfr), a male with complete male sterility were recovered. This male
sterile phenotype is expressed. This form of male sterility was maternally inherited and perfectly
sterility can arise spontaneously or bred through wide maintained by a cultivar ICPL 288. It was designated
hybridization. The complete CMS-based hybrid system as A2 cytoplasm source. Subsequently, Saxena and
involves three distinct genotypes - the male sterile Kumar (2003) also bred CMS lines by crossing C.
female (A-) line with ‘S’ cytoplasm and recessive scarabaeoides (as female) with cultivar ICPL 88039
fertility (frfr) nuclear alleles; the maintainer (B-) line (as male).
carrying fertile normal ‘N’ cytoplasm and the same
Breeding of another (A4) CMS system
recessive (frfr) nuclear alleles. The third parent,
designated as restore (R-) line, contains dominant (FrFr) Another stable source of CMS was bred at ICRISAT
fertility restoring nuclear alleles and it is responsible by crossing C. cajanifolius with cultivar ICP 28 (Saxena
for inducing male fertility in hybrid (A x R) plants. et al. 2005a). In this cross all the hybrid plants were
Further, depending on the number and type of fertility male sterile and it was maintained by backcrossing
restoring nuclear genes and specificity of their the hybrid plants with ICP 28. This CMS-inducing
interaction with mitochondrial genes, the expression cytoplasm, designated as A4 system, is being used
of male fertility/sterility in hybrid plants could be total in breeding hybrids at ICRISAT, ICAR and some public
or partial. Sometimes such expressions are also and private seed companies.
affected by prevailing environmental conditions such
as photo-period, temperature, or both. Fertilty restoration

In search of good fertility restorers of A4 CMS system,

Since extensive search in the pigeonpea
over 3000 testers were examined at ICRISAT and
germplasm did not yield any CMS system, elaborate
various ICAR centres. From the results it was
plans were made to breed this system by integrating
concluded that i) plenty of fertility restorers were
nuclear genome of the cultivated type with
available in the germplasm, ii) the fertility restoring
cytoplasmic genome of a wild relatives of pigeonpea.
genes were distributed randomly with greater frequency
The first unsuccessful attempt in this direction was
in the medium maturing germplasm, and iii) in early
made by Reddy and Faris (1981) when they crossed
maturity the fertility restoring genes were relatively
C. scarabaeoides and C. sericeus, the wild relatives
less frequent (Dalvi et al. 2008; Saxena et al. 2014
of pigeonpea, as female parent with a C. cajan line as
a,b). Some early maturing restorers, identified during
male parent. Although in BC1F2 generation, some
the study, could not produce stable hybrids with respect
segregants exhibited male sterility that was inherited
to their fertility restoration. A perusal of published
maternally, but unfortunately, these plants suffered
reports on the genetics of fertility restoration of A4
with female sterility and abnormal (petloid) flowers.
CMS system showed that either one or two dominant
Hence, all the selections were discarded.
genes controlled the fertility restoration in F1 generation
Ariyanayagam et al. (1995) launched a targeted
with variable gene action. In a significant study Saxena
breeding programme to develop CMS using
et al. (2011) reported that the fertility restoration was
mutagenesis and wide hybridization approaches.
stable across environments only when two dominant
Progenies of a cross between C. sericeus and a cultivar
genes were present together in a single hybrid; and
exhibited maternal inheritance for male sterility, but
the hybrid combinations carrying either of the dominant
for some reasons, this trait could not be stabilized for
genes were also fertile but not in all the environments.
use in hybrid breeding.
Phase III (2003-2013): process of releasing the
Success in breeding the first (A2) CMS system
world’s first commercial pigeonpea hybrid
The first workable CMS system in pigeonpea was
Release of the first CMS-based pigeonpea hybrid
produced at Gujarat Agricultural University, SK Nagar
GTH 1 in Gujarat
by Tikka et al. (1997). It was derived from a naturally
cross-pollinated (an off-type) plant observed in the The first CMS-based pigeonpea hybrid GTH-1 was
August, 2015] Hybrid pigeonpea review 285

developed at GAU, SK Nagar, and released by ICAR Table 3. Performance (yield kg/ha) of three hybrids in
in 2004 for cultivation in Gujarat state. This hybrid the on-farm trials
was bred by crossing A 2 CMS line GT 288A with
Hybrid State Farmers Hybrid Control Stan-
fertility restorer GTR-11. Based on yield trials
(no.) yield yield dard
conducted at multi-locations during 2000 to 2003, GTH hete-
1 (mean yield 1830 kg/ha) recorded 32% yield rosis (%)
superiority over the best local check (GT 100/101)
GTH-1 Gujarat 04 2673 1996 25
(mean yield 1330 kg/ha). This hybrid is non-determinate
in growth habit and early in maturity (140 days). Its ICPH 2671 Maharashtra 782 969 717 35
Andhra Pradesh 399 1411 907 56
flowers are yellow and seeds are large and white. In
Jharkhand 288 1460 864 69
front-line demonstrations conducted in three districts
Madhya Pradesh 360 1940 1326 46
in 2003, the hybrid recorded 25.3% yield advantage Total/mean 1829 1445 954 51
over popular control (Table 3). After multi-locational
ICPH 2740 Madhya Pradesh 13 1814 1217 49
evaluation by ICAR, the hybrid GTH-1 was released Andhra Pradesh 47 1999 1439 39
for cultivation in Central Zone. Unfortunately, in spite Gujarat 40 1633 1209 35
of its high performance in farmers’ fields, this hybrid Total/mean 100 1825 1288 41
could not be commercialized due to its inability to Source: various reports and publications of ICRISAT and ICAR
produce uniformly male fertile flowers and record good
yields in different locations/environments.
3). In Maharashtra state, a total of 782 on-farm trials
Release of the first commercial hybrid in pigeonpea were conducted in seven districts and the hybrid
Hybrid ICPH 2671 is the first ever commercial hybrid produced 35% more grain yield (969 kg/ha) than the
of any grain legume. It was produced by crossing a control. In Andhra Pradesh (399 trials), the hybrid
male sterile line ICPA 2043 with restorer ICPR 2671. exhibited 56% superiority over the control. Similarly
The plants of ICPH 2671 are semi-spreading, non- in Madhya Pradesh (360 trials), the hybrid out-yielded
determinate with profuse branching. It grows over two the control cultivar by 46%; while in Jharkhand (288
meter in height, matures between 164-184 days and trials) ICPH 2671 demonstrated 69% superiority over
contains 3.7-4.0 seeds/pod. The purple coloured seeds the control cultivar Bahar. Over all the four states (1829
weigh between10.5 to 11.2 g/100 seeds. ICPH 2671 trials), ICPH 2671 produced 1445 kg/ha average yield
has high level of resistance to both wilt and sterility and it was 51% more than the local checks (954 kg/
mosaic diseases. In comparison to inbred cultivars ha). ICPH 2671 was released for general cultivation in
the hybrid, by virtue of its greater root mass and depth, 2010 by the state Government of Madhya Pradesh
possesses greater ability to draw moisture from deeper and later notified for cultivation in the entire country
soil profiles. Its fast root growth also helps in (Saxena et al. 2013). After this release, two more
overcoming short spells of early season drought that hybrids ICPH 2740 in Andhra Pradesh (Saxena et al.
is often encountered in July-sown rainfed crops. ICPH 2014c) and ICPH 3762 in Odisha (Saxena et al. 2014d)
2671 also recorded have high survival (88%) under were also released for cultivation.
water-logged conditions; and this was found to be
Large-scale hybrid seed production of ICPH 2671
related to its ability to utilize stored assimilates through
anaerobic metabolism (Sultana et al. 2013). For hybrid pigeonpea seed production so far there are
no officially prescribed guidelines, but according to
During 2005-2008, ICPH 2671 was tested in the information generated by ICRISAT and various
multi-location trials and its mean performance in ICAR centres, an isolation distance of at least 500 m
different years ranged from 2200 to 3183 kg/ha; and is necessary. Since the extent of pollen transfer is
on average, it recorded 47% superiority over national determined by the population of pollinating insects,
check Maruti. In All India Advanced Hybrid Trials the row ratio of female to male may vary from 3:1 to
conducted at six locations in 2007, ICPH 2671 recorded 4:1. The rouging operation is recommended at seedling,
35% yield advantage over the control cultivar. In All flowering, and pre-harvesting stages. It is also advised
India Coordinated Trials, the hybrid (2564 kg/ha) that for optimizing hybrid yields, the adoption of proper
recorded 31% superiority over the control variety Maruti agronomy is essential (Mula et al. 2010 a, b). Our
(1996 kg/ha). In 2009 and 2010, the hybrid was experience has shown that the hybrid seed can be
evaluated in on-farm locations in four provinces (Table produced easily by growers, if the pollinating vectors
286 K. B. Saxena [Vol. 75, No. 3

are present in sufficient number. For other details

regarding hybrid pigeonpea seed production
technology, readers are advised to refer Saxena (2006).
In order to demonstrate the feasibility of large-scale
hybrid seed production, 94 professional seed producers
were selected in six states and the production
programme was undertaken under the direct
supervision of a team of experts. On average, 1019
kg/ha of hybrid seed was harvested and the highest
yield (1674 kg/ha) was recorded in Madhya Pradesh.
It is estimated that with a recommended seeding rate
of 5 kg/ha, an encouraging seed-to-seed ratio of 1: Fig. 2. Demonstration of seed purity assessment of
200 to 1:300 can be achieved. hybrid ICPH 2671 with the CcM 0021 marker.
The female lines, ICPA 2043 (298bp) and male
Seed quality control line ICPR 2671 (301 bp) show clear peaks using
a diagnostic SSR marker (CcM 0021, and the
In order to ensure consistency in performance of true hybrid (ICPH 2671) shows the presence of
hybrids in farmers’ fields, it is important that a high both the alleles (298 and 301 bp). Source :
level of genetic purity is maintained year-after-year. ICRISAT
In most field crops it is achieved by assessing the
quality of freshly harvested seed of hybrids and their
parents is determined by standard grow-out tests but identified by naked eye and called as “naked eye
in pigeonpea this approach is not feasible due to its polymorphic marker” is used to assess purity. Saxena
long generation turn-over time that may extend from 6 et al. (2011) identified ‘obcordate leaf’’ as a
to 9 months. To overcome this constraint two polymorphic marker and incorporated it in to A- and B-
approaches, briefly described here, have been lines. This marker, controlled by a single recessive
launched by ICRISAT. gene, can be easily recognized within a month from
sowing. The hybrids developed by crossing the parents
The first approach is based on genomics and it involving normal and obcordate leaf types will always
is simple, rapid, and cost effective. A set of 148 have normal leaves and the unwanted sibs will have
simple sequence repeat (SSR) markers was used for obcordate leaves. Such off-types can be detected
polymorphism survey of 159 hybrid parents and of within a month from sowing. This approach of hybrid
these, 41 markers were found polymorphic (Saxena breeding should be promoted to help in maintaining
et al. 2010). Subsequently, 3072 SSR markers were seed quality of female parents and hybrids.
also screened and a set of 42 SSR diagnostic markers
was identified for assessing the hybridity of ICPH 2671 Phase IV (2013-to date): pathways for producing
(Bohra et al. 2012). With the help of these markers the next generation hybrids
(Fig. 2) reliable detection of off-type hybrid seeds in
Breeding hybrids with specific adaptation
the commercial lots can be undertaken (Saxena et al.
2010; Bohra et al. 2012). Since at commercial level In modern agriculture the importance of specific
analysis of a large number of seed samples from adaptation of cultivars to a given environment cannot
different locations will be required, an alternative cost be ignored. Many farmers have now moved above the
effective genomics-based quality testing approach will subsistence level and regard agriculture as a
be an asset. To achieve this objective, several single challenging business. The prices of quality agricultural
nucleotide polymorphism (SNPs) have also been commodities are also high enough to attract greater
discovered for genotyping using Kbio Sciences Alleles investment for reaping more profits. The pigeonpea
Specific (KASPar) assays. This would allow assessing hybrids released so far have demonstrated high (2000-
genetic purity of large number of seed samples with 3000 kg/ha) yields and wide adaptation in farmers’
minimum number of loci. field conditions. Besides this fact, it was also observed
that some farmers harvested exceptionally high yields
The second approach involves breeding of hybrid
from the same hybrid in fairly large plots. For example,
parents with a naked-eye polymorphic marker: In this
under good crop management a group of progressive
approach a distinct phenotypic trait that can be
farmers in Maharashtra state harvested 4000-4500 kg/
August, 2015] Hybrid pigeonpea review 287

ha or more grains with >50% superiority over the through both resistant x resistant and resistant x
control. These observations emphasize the importance susceptible crosses.
of specific adaptation in breeding pigeonpea hybrids.
Saxena and Raina (2001) analyzed data from 11 Cytoplasmic diversification of A-lines
o o
diverse environments (10 -29 N) and reported The historical outbreak of southern corn leaf blight
significant effects of environments on the productivity disease in the USA (Tatum, 1971) gave a strong
of hybrids. They further concluded that greater yields natural lesson to breeders about the significance of
can be obtained by breeding locally adapted hybrids. cytoplasmic diversity in hybrid breeding programmes.
Chauhan et al. (1993) also advocated specificity of This happened because at one stage all the commercial
adaptation in early maturing cultivars. In this context, corn hybrids in the USA were based on ‘T cytoplasm’
it is not unfair to believe that soon the hybrid pigeonpea that carried genes for the susceptibility to blight
breeding programmes will grow in size and more disease; and when its outbreak occurred, most hybrids
specifically adapted hybrids will be available to provide succumbed and resulted in severe yield losses. In
more returns to farmers. this context, in pigeonpea a good beginning has already
Diversification of hybrid parents made and at present eight CMS-inducing cytoplasms
are known (Saxena et al. 2010; Saxena 2013).
Success of a dynamic hybrid research and Although these CMS systems represent a wide genetic
development programme, primarily depends on the variability for mitochondrial genome, so far only two
rate at which new parental lines are bred having high systems (A 2 and A 4 ) have been used in hybrid
combining ability and key market-driven traits. Such breeding. This situation necessitates breeding of more
parental lines can either be bred using both traditional number of CMS lines with greater cytoplasmic
and new technologies. In pigeonpea, the selection diversity. It should also be noted that while breeding
efficiency is adversely affected by out-crossing in the for cytoplasmic diversity, the effect of cytoplasm on
preceding generation. Hence, care should be taken to yield and other traits should also be studied. Recently,
minimize the incidence of natural hybridization in Saxena et al. (unpublished) reported some adverse
breeding plots. In the following text some strategies effects of C. cajanifolius (A4) cytoplasm on seed yield;
for diversifying the hybrid parents are discussed. but it was found to be cross-specific with the maximum
cytoplasmic-induced yield penalty of 19.5%.
Diversification of A-lines
There is a need to diversify A-lines with respect to Nuclear diversification of R-lines
both nuclear as well as extra-nuclear genomes. The Perfect fertility restoration and its stability are
available A-lines carrying A2 or A4 cytoplasm have important aspects in hybrid breeding programmes. In
perfect male-sterility system with high stability across early maturity group during the past few years, a total
diverse environments. To diversify the genetic of 46 fertility restorers have been identified among
backgrounds of these lines for use in hybrid breeding advanced breeding lines and germplasm (Table 4).
programmes, the male sterility was transferred into a These restorers exhibited >90% pollen fertility in hybrid
number of genotypes of diverse origin. In early group, combinations (data not reported) and have a wide range
the A4 male sterility has been transferred in 11 extra for flowering (50-85 days), maturity (101-141 days),
early and early maturing inbred lines through and seed size (6.2-12.1 g/100 seeds). None of the
backcrossing. The A 2 CMS system was also restorers had resistance to fusarium wilt, while ICPL
transferred to 19 diverse early maturing genotypes 149, ICPL 150 and ICPL 151 were resistant to sterility
(Parmar et al. 2008). The converted lines exhibited a mosaic disease. For A2 system also, a number of
wide range for flowering (60-81 days), maturity (105- fertility restoring lines were identified (Chauhan et al.
128 days), plant height (71-157 cm), and seed size 2004). The number of restorers in the early maturity
(8.1-12.5 g/100 seeds). In medium and long duration group is limited and more diversity would be required
group, >50 A-lines were bred at ICRISAT and various for a fruitful hybrid breeding programme. In medium
ICAR centres and many of them have resistance to maturity group, 113 fertility restorers were identified
both wilt and sterility mosaic diseases. In this context, with a considerable range for maturity, seed size and
the incorporation of dominant gene conferring most importantly disease resistance (Table 4). This
resistance to fusarium wilt (Saxena et al. 2012) into variability can sustain hybrid breeding for a few years;
A-lines should be given a serious attention because but breeders should look towards breeding more
this will allow development of wilt resistant hybrids
288 K. B. Saxena [Vol. 75, No. 3

Table 4. Variation for important traits among fertility most stable A-lines (ICPA 2039 and ICPA 2092) with
restorers of early, medium, and late maturity high general combining ability, were crossed with 502
groups recorded at Patancheru testers from the germplasm (Saxena et al. 2014a) and
among F 1 progenies, 179 were fully fertile, 26
Trait Early Medium Late maintained male sterility, and the remaining 297
(n= 46) (n= 113) (n= 31)
segregated for fertility and sterility. This study
Days to flower 50-85 90-130 131-158 suggested that the frequency of fertility restoring genes
Days to mature 101-141 138-200 186-241 in the germplasm is quite high and this resource can
Plant height (cm) 70-165 90-228 135-260 be exploited in hybrid breeding programmes. Overall,
this is the best short cut method for the diversification
100-seed wt (g) 6.2-12.1 6.8-17.3 7.7-18.1
of hybrid parents.
Wilt (%) 52-100 0-100 0-100
Sterility mosaic (%) 3-67 0-100 0-100 Formation and use of heterotic pools is another
proven approach for selecting hybrid parents, which
Source: ICRISAT
in high probability, produce high yielding hybrids. Shull
(1908) and Richey (1922) observed that the maize
diverse restorers. For late maturing group also, there hybrids involving diverse/dissimilar parents produced
is a need to search more restorers with diverse genetic plants with greater vigour and yield. Subsequently, for
backgrounds. Breeding new fertility restorers should discriminating the hybrid parents for their ability to
be a continuous process in a dynamic hybrid breeding produce high yielding hybrids, Sprague and Tatum
programme. To achieve this, besides normal pedigree (1942) evolved the popular concept of ‘combining
breeding, there are other approaches such as selection ability’. All these ideas of identifying good hybrid
from heterotic crosses and screening new germplasm parents gradually sunk into the concept of ‘heterotic
which can produce productive fertility restorers. groups’. This involves clustering of parental lines on
the basis of their performance in F 1 generation,
Greater productivity of hybrids is conditioned by combining ability, origin, phenotypic or genetic
various types of gene actions. The beneficial effects diversity. In pigeonpea, the first such information was
of genetic variation arising due to dominance, over published by Saxena and Sawargaonkar (2014) and
dominance and epistatic components will be lost in they constructed seven heterotic groups using specific
the subsequent self-generation; but its additive genetic combining ability effects from multi-location hybrid
fraction can be fixed through appropriate pedigree trials. According to this concept, crosses between the
selection. Saxena and Sharma (1990) and Sharma lines originating from the same cluster (group) are not
and Dwivedi (1995), while reviewing the subject, likely to produce heterotic hybrids; while the crosses
concluded that in pigeonpea additive genetic variation between the lines representing two diverse groups will
plays an important role in the formation of seed yield. have high probability to yield hybrids with high
This means that at least in some inbred lines the performance.
desirable alleles, present in the two parents, can be
brought together to realize high yields. These inbreds Search for new heterotic hybrids
can form good parental base for producing the new All the four released hybrids have recorded a
generation high yielding hybrids. A similar exercise satisfactory (25-50%) level of standard heterosis in
carried in hybrid ICPH 8 revealed that some inbred farmers’ fields. Considering the vast area and variable
lines achieved 70-75% yield potential of the hybrid agro-ecological conditions, there appears a need for
(Saxena et al. 1992). Use of such inbreds in hybrid breeding many more hybrids for the country. Efforts
breeding is expected to yield hybrids with productivity in this direction are being made both at ICRISAT and
greater than the available hybrid combinations. ICAR. In this context, several early maturing
The primary gene pool of pigeonpea with >15000 experimental hybrids were evaluated for their fertility
collection maintained at ICRISAT and ICAR is a rich restoration and productivity (Saxena et al. 2005b); and
source of genetic diversity for almost all the important eight promising combinations were advanced to multi-
quantitative and qualitative traits. This genetic wealth location testing (Table 5). From this material, two
can be exploited in identifying new maintainers and hybrids ICPH 2433 and ICPH 2438 were found
fertility restorers for use in hybrid breeding promising with respectively, 54% and 42% superiority
programmes. To start this activity at ICRISAT, two over the control UPAS 120. Recently ICPH 2438 has
August, 2015] Hybrid pigeonpea review 289

been identified for on-farm trials in Madhya Pradesh over the well-known ‘three line’ system. These include
(AK Tikle, personal communication). The performance i) multiplication of A- line without using the maintainer
data demonstrated that the level of hybrid vigour in line and pollinating agents, ii) elimination of fertility
early maturity pigeonpea group is high enough for restorers from hybrid system, iii) elimination of
commercialization. A perusal of productivity data also deleterious cytoplasmic effects, iv) utilization of greater
showed that in comparison to inbred cultivars (12.5 genetic variability in hybrid breeding, and v)
kg/ha/day), the hybrids were more efficient in dry matter development of a large number of hybrids in a short
production and/or it’s partitioning in to grains (22 kg/ time. In pigeonpea, the temperature sensitive male
ha/day). Further studies are needed to understand the sterility (TGMS) is of recent origin (Saxena, 2014) and
physiological aspects of yield determination in hybrids, it was derived from an inter-specific cross involving
their inbred parents, and control cultivars. Information C. sericeus and C. cajan. Plants carrying the sensitive
on the role of yield contributing traits needs to be gene are completely male sterile at the temperatures
generated for understanding the reasons for relatively
greater productivity of the hybrids.
Table 6. Some promising medium duration pigeonpea
In the last few years over 1500 medium duration hybrids developed at ICRISAT
hybrids representing maturity groups V and VI were
Hybrid Yield Standard 100-seed *Wilt *Sterility
developed; and based on yield and disease resistance
(ICPH) (kg/ha) heterosis weight (%) mosaic
eight combinations were selected for on-station testing. (%) (g) (%)
These hybrids exhibited 37-62% standard heterosis
and high levels of resistance to wilt and sterility mosaic 3371 3013 62 11.50 0 0
diseases (Table 6). Pandey et al. (2013) also recorded 3491 2919 57 13.40 0 0
over 20% standard heterosis in four long duration 3497 2686 44 10.90 0 15
pigeonpea hybrids. 3481 2637 41 11.60 0 0
Exploring temperature-sensitive male sterility in 3494 2586 39 12.40 0 9
hybrid breeding 2740 2900 57 12.30 0 0

Some environmental factors such as temperature and 3762 3000 62 11.90 0 0

photo-period are known to play a key role by inducing 2671 2509 37 12.20 0 0
male sterility and fertility in various crops (Kaul 1988). Check 1864 - 11.10 0 0
However, the credit of using such temperature and SEm ±205.7 - ± 0.33 - -
photo-period induced male sterility in breeding hybrid
Mean 2448.1 - 11.7 - -
rice goes to Chinese scientists (Shi 1981; Zhou et al.
1988; Sun et al. 1989). This approach, popularly known CV (%) 11.9 - 3.98 - -
as ‘two parent hybrid breeding’ has various advantages *Data from disease nursery

Table 5. Performance (yield kg/ha) of early maturing hybrids (A4) in multi-location trials

Hybrid Maturity 2007 2008 2009 2010 Mean % Prod.

(ICPH) (days) (n=7) (n=4) (n=8) (6) (n=25) gain (kg/ha/d)
2433 114 2538 1864 2331 2489 2306 54 22.22
2438 115 2722 1570 2238 1979 2127 42 18.50
2363 115 2292 1763 2131 2005 2048 36 17.81
2429 114 1825 1907 2015 2037 1946 30 17.07
2431 117 2186 1400 1925 2165 1919 28 16.40
2447 114 1959 1456 2045 1782 1811 21 15.89
2364 114 1909 1294 2018 1883 1776 18 15.58
3310 106 1540 1344 1731 1546 1540 3 14.53
Check 120 1502 1204 1545 1758 1502 - 12.52
Source: ICRISAT
290 K. B. Saxena [Vol. 75, No. 3

>25oC, while at <24oC the same plants become fully develop heterotic groups and for the selection of
male fertile. Field evaluation of this material at potential genotypes for breeding high yielding hybrids.
o
Patancheru (17 N) showed that in June sown crop,
the plants were male sterile in the month of September Conclusion
(high temperature). The same plants turned fully male The decades-old productivity stagnantion in pigeonpea
fertile in the month of November (low temperature); has now become a serious concern at national level,
and when the temperatures rose in the month of particularly in view of the national nutritional security.
February, they again reverted back to male sterility. To overcome this GMS-based technology GMS-based
To make full use of this system in pigeonpea ‘critical technology constraint partly, a CMS-based hybrid
fertility point (CFP)’ and ‘critical sterility point (CSP)’ pigeonpea breeding technology has now been evolved
that determine the conversion of male sterility/fertility and the hybrids have demonstrated clear advantages
need to be worked out. In India the two-parent hybrid over pure line cultivars for yield and stability. In this
system can be adopted easily as the sites with desired context, the release of three commercial pigeonpea
temperature requirements can be identified for seed hybrids is considered a major breakthrough in
production using different seasons, altitudes, and pigeonpea breeding. Extensive on-farm testing of the
latitudes. This system will require two sites each with hybrids in seven Indian states has shown that the
specific threshold temperature regime to allow full magnitude of standard heterosis in pigeonpea is
expression of the TGMS gene(s). For multiplication of comparable with other field crops, where commercial
female parent, the maximum safe mean temperature hybrids are already in market. The cultivation of
during reproductive phase at the production sites hybrids has also given positive signals to farmers
o
should not exceed 20 C; while the hybrid seed about the drought tolerance and high productivity of
production can be taken at most places in rainy season hybrids.
o
when the temperatures are well over 26 C. This trait
can be transferred easily to any inbred line (Saxena The hybrid seed production technology in
and Bharathi, 2015) provided the screening facilities pigeonpea is easy and high seed-to-seed ratio (1: 200
with controlled temperature are available. to 1: 300) ensures its economical production. However,
the selection of production sites and development of
Integration of genomics science in hybrid breeding a seed chain are essential forsustainable adoption of
Besides seed quality testing (Fig. 2), the genomics hybrid pigeonpea technology. Further, for the promotion
can also be used to identify fertility restorers without of hybrids, use of expert hands and minds will bring a
going into the cumbersome process of making hybrids big difference in the pace and quality of technology
and testing their progenies for the presence of genes transfer. In this context, training of seed producers
responsible for restoring their pollen fertility. The marker will be a step in the right direction. At present there
assisted breeding may help in the identification and are no prescribed seed standards for hybrids in
selection of lines carrying the Fr genes within pigeonpea. Such issues must be addressed at national
segregating populations and germplasm at a faster level and the information be made available to all the
pace and economically. This is facilitated by seed producers. Further, for large-scale adoption of
constructing a consensus genetic map using the hybrids, it is also necessary to convince both public
populations segregating for the fertility restoring genes. and private seed companies about the benefits and
The first pigeonpea consensus map has already been profitability of hybrid pigeonpea business.
constructed (Bohra et al. 2012) and it contains 339
The hybrid technology has now provided a new
simple sequence repeat (SSR) loci spanning a distance
instrument for realizing quantum jumps in the
of 1,059 cM. In three mapping populations a total of
productivity of pigeonpea; and with a conservative
four major quantitative trait loci (QTLs) for fertility
view, it is estimates that with 25-30% on-farm yield
restoration have been identified. This technology has
advantage and about 10% replacement of pigeonpea
a great potential and research on its refinement is
area with hybrids, India can produce enough pigeonpea
underway in the genomic laboratories at ICRISAT and
to meet its domestic needs.
ICAR. The genomics science can also be used to
generate accurate information on genetic diversity of Acknowledgements
hybrid parents. This will eliminate environmental effect
which may mask the true genetic variability in the traits The contribution of various ICAR research centres
of interest. This information can also be used to including PAU, Ludhiana; IARI, New Delhi; IIPR,
August, 2015] Hybrid pigeonpea review 291

Kanpur; PKV, Akola; GAU, SK Nagar; ARS, Sehore; Dalvi V. A., Saxena K. B. and Madrap A. I. 2008. Fertility
TNAU, Coimbatore; APAU, Hyderabad; MAU, restoration in cytoplasmic nuclear male sterile lines
Parbhani; ARS, Gulberga and Mahyco, Jalna in derived from three wild relatives of pigeonpea. J
Hered., 99: 671-673.
conducting joint evaluation of hybrids and sharing
research and development related materials and Green J. M., Sharma D., Saxena K. B., Reddy L. J. and
information is duly acknowledged. The financial Gupta S.C. 1979. Pigeonpea breeding at ICRISAT.
Paper presented at the Regional Workshop on
support received from National Food Security Mission,
Tropical Grain Legumes. 18-22 June 1979.
DAC. New Delhi is also acknowledged. Author would University of West Indies. Trinidad.
also like to thank colleagues and technicians at
Green J. M., Sharma D., Reddy L. J., Saxena K. B., Gupta
ICRISAT and other research centres for their full S.C., Jain K.C., Reddy B.V.S. and Rao M. R. 1981.
support. Methodology and progress in the ICRISAT
pigeonpea breeding program. In: Proc. Intl.
Workshop on Pigeonpeas. International Crops
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