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Journal of Biotechnology

& Crop Science Review

5(6): 32-40, 2016

Breeding hybrids and inbreds for temperature tolerance in maize

VP Mani, RS Khandelwal, GS Bist, KS Koranga
Received: 15 March 2016 Revised Accepted: 10 April 2016


The important breeding objective in any crop plants is to improve grain yield potential and stability of the cultivars including
in maize. Stability of performance becomes more important if crop is grown in less favorable environments. Tolerance to
abiotic stresses is in general and drought through cold and heat tolerance in particular is essential in crop plants, as plant
cannot alter an environmental stress external to itself. The effect and mechanism of all the three types of temperature stresses
vary with respect to survival, growth and development and physiological processes. So during planning of crop improvement
in open pollinated crops like maize for temperature tolerance through development of inbred and hybrids one must keep in
mind regarding mechanism of temperature tolerance. The improvements in vegetative growth and yield performance of any
of these crops will require selection of inbreds for hybrid development that have more efficient growth habits under
unfavourable stresses conditions.

Key Words: Drought, Hybrids, Inbreds, Maize, Resistance, Tolerance


Temperature stress tolerance of a plant is an ability of the field, become more resistant and therefore capable
the plant to come in thermodynamic equilibrium with of surviving chilling temperatures. However, in
extreme temperature fluctuations without suffering summer and rainy season the plant withstand higher
injury. Hybrids or Inbreds with heat and cold stress temperature stress at early vegetative phase and low-
tolerance is able to prevent, decrease or repair the temperature stress at reproductive phase. When the
injurious strain caused by the stress at a negligible tissues of the plant are frozen below a threshold
loss in the economic yield, provided proper and freezing temperature specific for that crop, it may
scientific methods are followed. The heat and cold cause an irreversible freezing damage to the plant.
stresses on plant have different susceptibility at The development of inbred and hybrids that
vegetative and reproductive phase. avoid/tolerate freezing/chilling temperatures should
be targets of a plant breeder to sustain and increase
In winter, the crop generally suffers cold stress at economic yield in crop plants under stress conditions.
germination especially in late sown conditions and Even though plants are categorized into six response
high temperature stress at reproductive phase during classes with respect to low temperatures (Levitt 1980)
grain filling period. During vegetative phase, the crop but for a plant breeder, these six categories can be
plants may face chilling or frosting damage. Crops in aggregated into two broad groups based on nature of
tropical and sub-tropical climates show varying stress, its physiological repercussions, the nature of
degrees of resistance to chilling injury. The plants tolerance and the agronomic implications. These
that are susceptible to chilling injury may harden in groups include plants, which are chilling sensitive
and hardy plants.The most important breeding
VP Mani, RS Khandelwal ( ), GS Bist, KS Koranga objective in crop plants is to upgrade grain yield
Vivekananda Parvatiya Krishi Anusandhan Sansthan (ICAR)
Almora 263601, Uttaranchal potential and stability of performance. Stability of
Email: performance is very important when crop is grown in

J of Biotech & Crop Sci (2016) 5(6): 32-40

less favourable environments. Tolerance to abiotic the phenotypic characters that could assist a breeder
stresses is essential in crop plants, as plant cannot in successful implementation of the programme.
alter an environmental stress external to itself.
Heat Tolerance, its Mechanism and Genetic Basis:
Temperature Tolerance System Temperature higher than the optimal is considered as
heat stress that may be at vegetative growth or at
It is important to understand the system of inheritance reproductive phase. At vegetative growth, the adverse
of temperature stress for any corp for its improvement effect of high temperature may be on seedling
and development of hybrids and inbred in crop like emergence, vigor and survival, while at reproductive
maize. Temperature stress includes chilling, freezing stage, the effects may be on process of fertilization,
and heat stress a crop has to withstand during its pollen viability, stigma and style, ovules and post-
vegetative as well as reproductive stage. The ability anthesis. At the whole plant level, heat stress causes
of the plant to recover from the consequences of a an acceleration of developmental stages, abortion of
heat stress is important for the crop yield and seed development, and impairment of grain filling. At
stability. Manipulation of regulatory processes that the physiological level, high temperatures affect
are responsive to oxidase stress could enhance major functions of plant cell, including
chilling tolerance mechanism, while a precise model photosynthesis, energy metabolism, and translocation
for freezing tolerance is yet to be determined. The of assimilates. Plant enzyme activities are
genetic basis of inheritance of winter hardiness is thermolabile, including RUBISCO (Weis 1981),
complex and polygenic in nature. Primarily, tolerance sucrose synthase (Rijven 1986), Catalase (Willekens
to temperature stresses is largely non-additive. et al 1995) and superoxide dismutase (SOD) (Matters
However, the selection for SCA would allow the and Scandalios 1986). Inhibition of these enzymes is
recombination of temperature tolerance with other major cause of inactivation of CO2 fixation at high
agronomically desirable attributes. The effect and temperatures. Heat stress severely inhibits plant
mechanism of all the three types of temperature capacity to convert sucrose into starch. At molecular
stresses vary with respect to survival, growth and level, the most prominent alterations caused by heat
development and physiological processes. So during stress in plant cell are modification of protein
planning of crop improvement in open pollinated synthesis, including synthesis of Heat Shocked
crops for temperature tolerance through development Proteins (HSPs) and antioxidant enzymes, and an
of inbred, scientist must keep in mind regarding alteration of membrane composition and structure.
mechanism of temperature tolerance for all these Genetic variation in heat tolerance for various
types. Many of the important crops around the world attributes exist in crop plants viz. germination,
are now cultivated in areas where temperature stress growth during heat stress, growth recovery after heat
is common. Therefore, improvements in vegetative stress, photosynthesis, translocation, flowering and
growth and yield performance of any of these crops seed set and stability of the cellular membrane. In a
will require selection of inbreds for hybrid 6 x 6 diallel study in sweet corn, Williams et al
development that have more efficient growth habits (1969) found that inheritance of recovery from a 6-hr
under unfavourable stresses conditions (Greaves heat stock at 52OC behaved in partially dominant
1996). Before proceeding for developing inbred lines nature. Inbred showing a high GCA for the trait
or strains it is essential for a plant breeder to tended to produce more heat tolerant hybrids. In
understand the mechanism of temperature stress, another study on tomato, it was found that percent
genetics of response of crop plant to different stress normal flower and percent fruit set showed partial
conditions. While planning a breeding programme for dominance. Flower production and fruit set under
temperature stress tolerance, it is essential to identify heat stress appear to be genetically independent

J of Biotech & Crop Sci (2016) 5(6): 32-40

(Hanna et al 1983). Strong association between heat et al 1994). Since low temperature induces oxidative
tolerance and simple plant character cannot be stress in tissues, chilling damage is partly attributed
expected unless a genetic linkage is involved. Since to the effects of in-vivo-generated Reactive Oxygen
heat tolerance is largely a cellular manifestation, Species (ROS) in the cells. Kazemitabar et al (2003)
selection in cell cultures presumably carries and studied short term cold stress on rice seedlings and
potential as a method for improving heat tolerance. found alteration to the total leaf protein, including
Genetic resources for heat tolerance are apparently fragmentation of ribulose-1, 5-
quite common within the normal breeding germplasm bisphosphatecarboxylase RUBISCO). Physical
of different crops. On the other hand, for a good transitions of membranes from a liquid-crystalline
number of crop plants, the extent of existing genetic phase to a solid-gel phase were first suggested to be
variation in heat tolerance has a hardly been explored. the primary response of chilling-sensitive plants
The various studies on the genetic variation in heat subjected to low temperature stress (Lyons and
tolerance reveal a number of genetic resources for Raison 1970). Change in phase transition results in
heat tolerance in several important crop plants. increased leakiness of the membranes that ultimately
leads to a loss of campartmentation, collapsed
Chilling Stress, its Mechanism and Genetic Basis: gradient and disrupted metabolism (Levitt 1980,
Reduction in crop productivity by low temperatures, Wang 1982, Markhast 1986, Collings et al 1995).
called as chilling stress, whereby plants are exposed Chilling tolerance was also linked with the extent of
to low temperatures essentially above 0 OC. Chilling lipid unsaturation in some plant species (Roughan,
sensitive plants are typically tropical plants. Chilling 1985, Hugly and Somerville 1992). Efforts to alter
stress may be either at plant/organ level or at the sub- the degree of unsaturation of membrane lipids have
cellular level. At plant level, chilling stress affects resulted in improved chilling tolerance (Murata et al
seed germination, growth, fruit or seed set, economic 1992, Kodama et al 1994, Moon et al 1995). Levels
yield, pollen fertility and fruit quality. It leads to of some of the nuclear encoded proteins, such as F 1-
reduced germination, poor seedling establishment, ATPase, Cyt-oxidase, and molecular chaperones
stunted growth, wilting, chlorosis, necrosis, fruit or (HSP 50 and HSP 60) were also reduced in the
seed set, pollen sterility etc. while at sub-cellular mitochondria during chilling stress. The competence
level chilling affects membrane stability, chlorophyll and stability of mitochondria are important for plant
synthesis, photosynthesis, respiration and may tissues to survive low temperatures especially during
generate toxicity due to H2O2. In crop plants, chilling early seedling growth. Because mitochondria are both
injury results in poor seedling establishment, stunted the source and a target of oxidative stress in seedling
growth, wilting, chlorosis, necrosis, poor fruit set, etc. subjected to chilling (Purtarulo et al 1991, Prasad et
Water stress is often observed in chill affected plants al 1994, Purvis 1997), the functions of mitochondrial
and is largely localized at the root, as chilling of the proteins could be affected by chilling induced
intact root results in wilting (Berry and Raison 1981, oxidative stress. Chilling causes a decline in the
McWilliam 1983, Christiansen and John 1981). activity of cyt-oxidase and an increase in alternative
oxidase activity (Elthon et al 1986, Rybka 1989,
Some of the changes related to low temperature stress Prasad et al 1994). Genetic variation in chilling
include alterations in gene expression, composition of tolerance has been revealed for various physiological
proteins, lipids, carbohydrate, membrane properties, process such as photosynthesis and plasma-
solute leakage, mitochondrial respiration and membrane function as well as different plant
photosynthesis (Levitt 1980, Wang 1982, Markhart functions such as generation, growth seed set etc.
1986, Guy 1990, Thomashow 1990, McKersie 1991, However, most of the information is available on
Howarth and Ougham 1993, Smiranoff 1993, Foyer traits like seed germination and seedling

J of Biotech & Crop Sci (2016) 5(6): 32-40

establishment under low temperature. Field positively associated with seedling vigour (Stamp
emergence of maize under early planting was 1984). A positive association may therefore, exists
improved by recurrent selection at the rate of 84% between cold germination and seedling vigour,
(Mock and Bakri 1976). In a maize population, depending on the nature of the genetic material and
derived from a diallel mating system, primarily the environmental conditions under which selection is
additive genetic effects with a significant maternal perform. The question whether the association found
effect conditioned the emergence rate (Chapman are genetic or physiological is unresolved. The
1984). The large genotype x environment interaction prevalence of dominance gene action for various
revealed for emergence rate. In another study, attributes of chilling tolerance in plants is manifested
multiplicity of environmental factor that affects field and used in hybrids. Genetic resources for chilling
emergence of maize was reported (Unander 1984). tolerance are to be found in the lower temperature
margins of the crops range of distribution. High
In a study of 56 reciprocal single crosses and parental altitude races for various crop species were found to
lines of maize, the nature of inheritance of cold possess relatively high levels of chilling tolerance.
germination at 6OC and 8OC was found to be complex While ample genetic variation for chilling tolerance
with strong maternal effects (Pesev 1970). A possible has been identified in well-adapted breeding
genotype x temperature in most likely at the basis of population, such as the case is for maize (mock and
conclusion made that selection for chilling tolerance Eberhart 1972), the use of exotic tolerance races of
at germination must be performed at several low the crop has been shown to offer good opportunities
temperatures. When selection was performed in for improving its chilling tolerance (Mendoza 1979).
maize under different germination temperatures both The use of the wild relatives of crops has also been
additive and dominance variances were found to attempted. Introgression of chilling tolerance from
control the trait (Keim and Gardner 1984). Maternal such exotic germplasm has been successful in some
effect could not be ignored and the conclusion was cases, such as the tomato.
that selection under growth chamber condition did
not result in significant improvement of cold Freezing Tolerance, its Mechanism and Genetic
tolerance at germination. Basis: The freezing causes 70 to 80 per cent of the
liquid water in the tissues are frozen as extra cellular
Crop establishment under condition of early planting ice, resulting in cellular dehydration (Levitt 1980).
also involves chilling tolerance of the growing The abilities of organisms to modify the growth of ice
seedlings. The question is raised whether cold and to withstand extensive dehydration are key
tolerance at germination is associated either elements of cellular survival. Freezing may cause
genetically or physiologically with cold tolerance of disruption in and/or alter the semi-permeable
the growing seedlings. In selection experiment properties of plasma membrane. This results in a loss
performed with adapted maize population, the rate of of solutes from the cells. Further, cells remain
cold emergence was not consistently or positively plasmolyzed even after thawing. In addition, cells
correlated with seedling dry matter. Selection for cold with intact plasma lemma may take up excess water
germination in the growth chamber resulted in only and become unusually turgid. Thus, the freeze-thaw
little improvement in seedling vigour in maize. On cycle may create stress by affecting the structure and
the other hand, significant positive genetic correlation function of plasma lemma, that depends on the
was revealed between seedling dry weight and vigour stability of protein and lipid components under
and the total number of seedling emerged in the cold freezing stress. There are various components of
(Keim and Gardner 1984). In nine tropical maize freezing tolerance like osmotic adjustment, amount of
populations, the rate of cold emergence was bound water, plasma membrane stability, cell wall

J of Biotech & Crop Sci (2016) 5(6): 32-40

components and cold responsive proteins. For many species geographical distribution. While very hardy
plants like Brassica cold acclimation is associated materials may be revealed within the cultivated
with accumulation of soluble sugars in the cytosol, germplasm the available information shows that
especially sucrose (Fu et al 1999). transgressive segregation from hardy x hardy crosses
is rare. Genetic variation for freezing tolerance may
Numerous studies were devoted to the inheritance of also be exists in non-hardy materials. Induced
winter hardiness in the field, and conclusions were mutations are another source of genetic variability for
drawn with respect to selection work. Complexity of freezing tolerance. The wild relatives of crops were
GxE interaction, leading to the low observed often considered as genetic resource for freezing
heritability, was found. Marshall (1982) reviewed tolerance. The use of cell and tissues in selecting for
some of the results on heritability in winter cereals. freezing tolerance is practically non-existent. The
He noted studies where high heritability was revealed complexity of plant freezing resistance would seem to
up to 93%, especially after years of consistent and hinder any practical application of in vitro selection
differential winterkill. He concluded that the high methods.
estimates are unduly encouraging because they do not
reflect the whole spectrum of possible stresses that HYBRID AND INBRED DEVELOPMENT
occur in the field. Grafius (1981) concluded that the IN MAIZE
inheritance of winter hardiness is complex and
polygenic in nature. Grafius (1981) noted for his own The pedigree method is most widely used to develop
work that from one year to the next year, the same inbred lines and consequently for hybrid development
set of varieties has exhibited a correlation of 0.60 in maize. The fixation of genes through inbreeding
with regards to winter survival. Therefore, winter can best be achieved through selfing followed by full-
hardiness, as a phenomenon measured in the field sib mating and half-sib mating. Repeated selfing
environment, is apparently not a genetic entity. helps in unmarking deleterious recessive genes,
which are selected against and advantageous genes
Freezing tolerance in the cereals under controlled are retained. Selfing results in uniformity and loss of
freezing conditions was largely controlled by additive vigour, therefore, it is important to select and test in
(Sukla 1981) or largely by non-additive (Parodi et al. each selfing cycle. One generation of self pollination
1983), gene action. When actual improvement of results in same level of inbreeding as three
winter hardiness is sought, enough genetic variation generations of full sib mating and six generation of
did not exist. Transgressive segregation for the trait half-sib mating (Hallauer and Miranda 1988). A
has to be revealed. In contrast, Limin and Flower slower approach of fixation of genes would be more
(1982) noted additive gene action with a few genes to useful in development of inbred lines for characters
control freezing tolerance in their wheat crosses. like temperature tolerance which are controlled by
Marshall (1982) discussed transgressive segregation polygenes thereby provide more opportunity for
for freezing tolerance in the cereals and its utilization selection. During phenotypic selection under stress
in the development of hardy varieties. Hard evidence environments, selections may be carried out both
on genetic association between freezing resistance between and within progenies. As the inbreeding
and to the plant characters is limited, inspite of the continues, the differences between progeny increase
fact that such association are of great importance. The and those within progenies decrease. At initial stages,
immediate and preferred germplasm would naturally more plants need to be selected giving more
be any cultivated forms that survive extreme winter opportunity for the desired genotype to progress
conditions. Such materials are likely to be found in further for final selection at later generations. Each
the extreme low temperature margins of the crops or generation need to be adequately sampled to include a

J of Biotech & Crop Sci (2016) 5(6): 32-40

range of segregants. For two parents differing for 10 development and maintenance of inbred lines through
allelic pairs, the F2 population size is 1,048,576 recurrent selection is the most effective method to
individuals. Hence with large number of genes improve for abiotic stresses. Inbred lines are
involved in the temperature stress tolerance, the improved by introgression of specific gene(s) from
sample size becomes very large. Practically such identified donor germplasm for desired agronomic
large populations are difficult to handle. Those cases characters. These genes can be available in distant
where a number of sources for temperature stress genera or species. The incompatibility barrier in
tolerance are available, it is practically preferred to transfer of such genes from distant species/genera can
have a smaller sample size and handle a large number be overcome using cell and tissue culture techniques.
of cross combinations. Bauman (1981) reported that Development of genetically engineered plants
500 were the most common sample size. Dhillon et al through genetic transformation holds great promise.
(1994) proposed the sampling of foundation plants The transgenic inbred line with insecticidal Bt gene is
for inbreeding, as half-sibs rather than self. This an example of commercialization of transgenic
approach nearly quadruples the effective population technology.
size and has other advantages. Tolerance to heat in
sunflower is an important abiotic stress in various The development of maize inbred line carrying a
ecological zones that contributed significantly in synthetic gene encoding a truncated version of Cry
increasing seed and oil yields. Selection of self- IA(b) derived from Bacillus thuringlensis for insect
fertility is also important in addition to heat tolerance resistance is an another example, expressed a high
among lines during inbreeding. level of insecticidal protein and showed resistance to
repeated infestation of corn borer in field conditions
Back crossing method works best for highly heritable (Koziel et al 1993). A number of successful efforts
characters and can make a good inbred even better. have been done in development of transgenic inbred
However, for a complex trait the results are variable lines in cotton, maize, groundnut, tobacco, tomato,
(Duvick 1974, Geadelmann and Peterson 1978). The potato, cucumber against various biotic stresses at
success of this method depends upon number of near-commercial level. However very little
genes controlling the inheritance of temperature information is available on development of transgenic
tolerance trait and the effect of environment on inbred lines carrying genes with tolerance to
selection. The numbers of back crossing vary cold/heat at vegetative and/or flowering stage.
depending upon recovery of the characters of
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