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Yield Improvement in Temperate Maize is Attributable to Greater Stress

Tolerance

Article  in  Crop Science · November 1999

DOI: 10.2135/cropsci1999.3961597x

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 Yield Improvement in Temperate Maize is Attributable to Greater Stress Tolerance
M. Tollenaar* and J. Wu
ABSTRACT
A retrospective analysis of the physiological basis of genetic yield
improvement may provide an understanding of yield potential and
may indicate avenues for future yield improvement. Rate of yield
improvement of maize (Zea mays L.) hybrids in Ontario, Canada has
been ≈ 1.5% yr21 during the last five decades. Comparison of short-
season hybrids representing yield improvement from the late 1950s
to the late 1980s showed that genetic yield improvement was 2.5%
per year and that most of the genetic yield improvement could be
attributed to increased stress tolerance. Differences in stress tolerance
between older and more recent hybrids have been shown for high
plant population density, weed interference, low night temperatures
during the grain-filling period, low soil moisture, low soil N, and a
number of herbicides. Yield improvement is the result of more effi-
cient capture and use of resources, and the improved efficiency in
resource capture and use of newer hybrids is frequently only evident
under stress. Improved resource capture has resulted from increased
interception of seasonal incident radiation and greater uptake of nutri-
ents and water. The improved resource capture is associated with
increased leaf longevity, a more active root system, and a higher ratio
of assimilate supply by the leaf canopy (source) and assimilate demand
by the grain (sink) during the grain-filling period. Improvements of
resource use under optimum conditions have been small, as leaf photo-
synthesis, leaf-angle distribution of the canopy, grain chemical compo-
sition, and the proportion of dry matter allocated to the grain at
maturity (i.e., harvest index) have remained virtually constant. Ge-
netic improvement of maize has been accompanied by a decrease in
plant-to-plant variability. Results of our studies indicate that increased
stress tolerance is associated with lower plant-to-plant variability and
that increased plant-to-plant variability results in lower stress tol-
erance.
A verage maize grain yield per hectare in Ontario
has increased at a rate of approximately 1.5% yr21
during the five-decade period since the 1940s, whereas
little or no improvement in grain yield was evident dur-
ing the five decades before the introduction of hybrids
in the 1940s (Fig. 1). Yield improvements of a similar
magnitude have been recorded for maize hybrids in
the USA and Europe (Tollenaar et al., 1994). Yield
improvement generally can be attributed to genetic im-
provement, changes in cultural management, climate
change, and the interactions among these factors. The
genetic improvement can be estimated from side-by-
side comparisons of hybrids that are representative of
the period under study. Estimates of the contribution
of genetic improvement to the overall yield improve-
ment in maize range from 40 to 100% (e.g., Derieux et
al., 1987; Tollenaar, 1989; Duvick, 1992). However, the
relative contributions of genetic gain and of gains due
to agronomic and environmental influences are difficult
Dept. of Plant Agriculture, Crop Science Division, University of
Guelph, Guelph, ON, N1G 2W1, Canada. Received 28 Dec. 1998.
*Corresponding author (ttollena@plant.uoguelph.ca).
Published in Crop Sci. 39:1597–1604 (1999).
1597
to separate, as genotype 3 environment interaction is
a prominent feature of yield improvement in maize.
Genotype 3 environment interaction effects on grain
yield are almost always significant for comparisons of
older and newer maize hybrids evaluated under two or
more different environmental conditions. Duvick (1984)
showed that differences in grain yield between older
and newer hybrids were a function of plant population
density. Differences between hybrids were small at low
plant population density and became greater as plant
population density increased. We have shown that a
more recent hybrid was influenced less by weed interfer-
ence than an older hybrid in a 5-yr field study conducted
in Ontario (Tollenaar et al., 1997). In contrast, differ-
ences in grain yield between the two hybrids were not
significant when the two hybrids were grown under opti-
mal belowground conditions (i.e., hydroponically) in the
field (Nissanka, 1995). Results indicate that a large pro-
portion of yield improvement in maize may be attribut-
able to the capacity of newer hybrids to better tolerate
stress conditions.
The observation that recent single-cross hybrids are
more stress tolerant than open-pollinated varieties and
older hybrids contradicts the notion that genotypic and
phenotypic variability are positively associated with
yield stability. Commercial maize genotypes in the U.S.
Corn Belt evolved from open-pollinated varieties before
the 1930s to double-cross hybrids from the 1930s to the
1960s and single-cross hybrids since the late 1960s. The
evolution of open-pollinated varieties of the 1930s to
single-cross hybrids of the 1990s was associated with an
increase in rate of yield improvement and an increase
in area of adaptation (Troyer, 1996). This evolution was
also associated with a decrease in genetic variability
within and among commercial maize genotypes: hun-
dreds of open-pollinated varieties were grown up to the
1930s in the U.S. Corn Belt, but the parentage of virtu-
ally all relatively recent, commercial U.S. hybrids in-
volve only six inbred lines or their close relatives (Good-
man, 1990). The notion that variability within a
community of plants, including mixtures of cultivars,
increases yield stability (Wolfe, 1985) suggests that yield
stability of an open-pollinated variety should be greater
than that of a single-cross maize hybrid. However, the
opposite effect is indicated by the increase in area of
adaptation and the increase in stress tolerance in the
comparison of older and newer commercial maize varie-
ties and hybrids. We have found that increases in yield
and stress tolerance of more recent maize hybrids grown
in Ontario are associated with enhanced crop stand uni-
formity (Wu, 1998).
Abbreviations: CV, coefficient of variability; PPFD, photosynthetic
photon flux density.
1598 CROP SCIENCE, VOL. 39, NOVEMBER–DECEMBER 1999
Fig. 1. Mean maize grain yield (0% moisture) in Ontario from 1892
to 1997, from Agriculture Statistics for Ontario, Ontario Ministry
of Agriculture, Food and Rural Affairs, Queen’s Printer of Ontario,
Toronto, ON, Canada.
We will analyze the physiological basis of genetic
improvement in maize in general and specifically for
short-season maize hybrids grown in Ontario. Yield im-
provement is the result of more efficient capture and
use of resources, and factors involved in the improved
efficiency will be reviewed. Second, the effect of stress
tolerance and stand uniformity will be discussed because
improved efficiency in resource capture and utilization
of newer hybrids is frequently only evident under stress.
Finally, we will speculate about the potential for genetic
improvement of maize in the future.
Resource Capture and Use
Resource Capture
Resource capture entails interception of incident so-
lar radiation by the maize canopy and uptake of nutri-
ents and water by the roots. Seasonal interception of
incident solar radiation, or photosynthetic photon flux
density (PPFD), can be enhanced by extending the pe-
riod during which 95% or more of incident solar radia-
tion is intercepted, which can be achieved either by early
canopy closure, delayed leaf senescence, or both.
Canopy Closure. Whereas enhanced seedling vigor
advances time of canopy closure and increases seasonal
PPFD interception, our results with short-season maize
hybrids show that total aboveground dry matter before
the 12-leaf stage of a newer hybrid was less than that
of an older hybrid in growth-cabinet studies (Tollenaar
et al., 1991; McCullough et al., 1994), and aboveground
dry matter of newer hybrids was lower than that of
older hybrids in a field study that involved six hybrids
(Tollenaar, 1996, unpublished data). Results of the
growth-cabinet studies indicated that lower early vigor
was associated with a greater root/shoot ratio. Root/
shoot ratio can influence rate of dry matter accumula-
tion during early development, because leaf area expan-
sion is directly related to amount of dry matter parti-
tioned to the leaves. Leaf photosynthetic rate did not
differ between the two hybrids (Tollenaar et al., 1991;
McCullough et al., 1994) and, consequently, plant dry
matter accumulation and rate of leaf area expansion
during early development were inversely related to the
root/shoot ratio.
Senescence. Delayed leaf senescence or “stay green”
is associated with yield improvement of maize hybrids
in North America (e.g., Crosbie, 1982; Tollenaar, 1991;
Duvick, 1997). Most of the differences in dry matter
accumulation between older and newer hybrids can be
attributed to the grain-filling period (Tollenaar, 1991;
Tollenaar and Aguilera, 1992). The “stay-green” charac-
teristic of the newer hybrid compared with an older
hybrid was associated with a larger source/sink ratio
during the grain-filling period (Rajcan and Tollenaar,
1999a).
Nutrient and Water Uptake. Nutrient uptake is re-
lated to root mass and energy supply. Limited data on
root mass during later phases of maize development
have been reported in the literature. A comparison of
the older hybrid Pride 5 with the newer hybrid Pioneer
3902 grown in a hydroponic system (Tollenaar and Mi-
gus, 1984) in the field in 1992 and 1993 showed that the
root/shoot ratio was ≈20% greater in the newer than in
the older hybrid during the grain-filling period (Nis-
sanka, 1995). Higher source/sink ratio in the newer hy-
brid (Rajcan and Tollenaar, 1999a) may imply that as-
similate supply to the roots is greater in the newer
hybrid. Rajcan and Tollenaar (1999b) reported that the
proportion of grain N derived from postsilking N uptake
was 60% for the newer hybrid and 40% for the older
hybrid. Continuous N uptake during the grain-filling
period has been associated with the ability to maintain
root growth after silking (Mackay and Barber, 1986),
which may be a function of assimilate supply. Water
uptake also may be related to root mass. Nissanka et
al. (1997) showed that decline in plant photosynthesis
during a drying cycle occurred on average 1 d earlier
in the older hybrid compared with the newer hybrid,
and the recovery of canopy photosynthesis after rewa-
tering was greater in the newer than in the older hybrid.
Resource Use
There are four avenues for increasing resource utiliza-
tion: an increase in gross leaf photosynthetic rate, an
increase in gross canopy photosynthetic rate, a reduction
in plant respiration, and an increase in harvest index
(i.e., grain yield as a proportion of total aboveground
dry matter at maturity).
Gross Leaf Photosynthetic Rate. An increase in gross
leaf photosynthesis may be the result of either an in-
crease of photosynthesis at saturating PPFD (i.e., Pmax),
an increase of photosynthesis at low PPFD (i.e., quan-
tum yield), or both. Differences in leaf net photosynthe-
sis between older and newer hybrids were not significant
under optimal conditions (Crosbie, 1982; Dwyer and
Tollenaar, 1989).
Gross Canopy Photosynthetic Rate. A more uniform
distribution of incident solar radiation across the crop
canopy can result in an increase in gross canopy photo-
 TOLLENAAR & WU: YIELD IMPROVEMENT AND GREATER STRESS TOLERANCE IN MAIZE
synthetic rate (Tollenaar and Dwyer, 1998). An increase
in leaf angle results in a more uniform distribution of
solar radiation across the canopy, and an increase in
leaf angle has been reported for Corn Belt hybrids be-
tween 1930 and 1990 (Crosbie, 1982; Duvick, 1997).
Effect of leaf angle on canopy photosynthesis can be
estimated from the change in PPFD distribution across
the leaf canopy and the photosynthesis-PPFD response
curve. Our estimates indicate a yield improvement in
the order of 20% when leaf angle increases from 30 to
608 (Tollenaar and Dwyer, 1998). In contrast with maize
hybrids in the U.S. Corn Belt, canopy architecture of
short-season hybrids in Central Ontario had not
changed by the late 1980s (Tollenaar and Aguilera,
1992), although leaf angle seems to have increased in
hybrids introduced during the 1990s (Tollenaar, 1996,
unpublished data).
Respiration. The two major components of respira-
tion are growth respiration, which is a function of the
composition of the plant dry matter, and maintenance
respiration, which is a function of the energy required
to maintain plant function and structure (Penning de
Vries et al., 1974, 1975). Little has been reported on
changes in plant composition associated with yield im-
provement in maize. Vyn and Tollenaar (1998) reported
that changes in grain composition associated with yield
improvement in Ontario were minor and Duvick (1997)
reported that grain protein concentration of U.S Corn
Belt hybrids declined by 1.8% yr21 from the 1930s to
the 1990s. A decline in protein concentration will result
in a reduction of growth respiration per unit grain, but
no data have been reported on total plant respiration
and its components. Earl and Tollenaar (1998) reported
a strong negative correlation between mature-leaf respi-
ration during the growing season and total seasonal dry
matter accumulation among three older and three newer
Ontario maize hybrids.
Harvest Index. The different responses of grain yield
to plant population density among maize hybrids repre-
senting various eras of breeding have led some to con-
clude that harvest index has been positively associated
with genetic improvement in maize (Russell, 1985). Har-
vest index declines when plant population density is
increased beyond the optimum plant population density
for grain yield in maize, and the optimum plant popula-
tion density for grain yield is lower for older hybrids than
for newer hybrids, which may explain the association
between harvest index and era of release for hybrids
grown at high plant population density (Tollenaar et
al., 1994). Harvest index did not differ among hybrids
when maize hybrids representing three decades of yield
improvement in Ontario were adjusted for optimum
plant population density (Tollenaar, 1989).
The Role of Stress Tolerance and Stand
Uniformity in Resource Use
Stress Tolerance
We define stress as a factor that causes, through either
its presence or its absence, a reduction in plant grain
yield. Consequently, stress is defined in terms of the
plant’s response to a causal factor. Stress can be allevi-
1599
ated either by management practices that influence the
causal factor or by modifying the plant such that the
impact of the causal factor on plant grain yield is re-
duced. Effects of stress in a production environment
(i.e., yields .2–3 Mg ha21) may be quite different from
those in a survival environment (i.e., yields ,2–3 Mg
ha21). Frey (1971) cited results of experiments con-
ducted in Colorado in which an open-pollinated variety
yielded 40% less than a maize hybrid in a production
environment with irrigation (i.e., 8.0 vs. 13.3 Mg ha21),
whereas yields did not differ under rainfed conditions
(i.e., 2.1 Mg ha21 for each of the two entries). Our
discussion on effects of stress on grain yield will be
limited to maize grown in a production environment
(i.e., yields .2–3 Mg ha21). Stresses that can influence
plant grain yield in a production environment are low
soil moisture, low soil N, low absorbed PPFD, and low
or high air temperatures. An increase in either the num-
ber of maize plants per unit area (i.e., plant population
density) or the number and size of weeds within a maize
stand will enhance the competition among plants for
resources within the maize canopy and, consequently,
stresses of this nature are termed compound stresses.
Grain-yield per unit area of a maize crop shows a curvi-
linear response to plant population density, with a maxi-
mum yield at the optimum plant population density.
Grain yield per unit area increases with increasing plant
population density until the increase in yield attribut-
able to the addition of plants is not greater than the
decline in mean yield per plant and, consequently, high
yield per unit area and plant stress are inevitably linked.
Results of our studies with short-season hybrids rep-
resenting grain-yield improvement in Central Ontario
have shown that newer hybrids are more tolerant to
stress than older hybrids. The differential response to
stress between older and newer hybrids has been shown
for low night temperature during the grain-filling period
(Dwyer and Tollenaar, 1989), low soil moisture in the
field (Dwyer et al., 1992), and under controlled-environ-
ment conditions (Nissanka et al., 1997), low soil N
(McCullough et al., 1994), and the herbicide bromoxynil
(3,5-dibromo-4-hydroxybenzonitrile, a Photosystem II
inhibitor) (Tollenaar and Mihajlovic, 1991). We also
have shown that tolerance of compound stresses such
as plant population density (Tollenaar, 1992) and weed
interference (Tollenaar et al., 1997) is greater in newer
than in older hybrids.
It has been suggested that selection under high plant
population density has been a key component in grain
yield improvement in corn (Troyer and Rosenbrook,
1983) and, consequently, changes in stress tolerance may
have been the result of indirect rather than direct re-
sponses to selection. One could speculate that such indi-
rect responses resulted from either or both (i) progres-
sively higher plant population densities in yield testing
programs and (ii) wide area testing , which has enabled
the evaluation of hybrids under a wide range of environ-
mental conditions.
Stand Uniformity
Although stand uniformity of field crops has long
been recognized as an important aspect of high-yielding
1600 CROP SCIENCE, VOL. 39, NOVEMBER–DECEMBER 1999
cultivars and breeders have been pursuing it since the
beginning of modern agriculture, it also has frequently
been claimed that cultivar mixtures can give higher and
more stable yields than pure lines in cereal crops (Wolfe,
1985). However, Marshall and Brown (1973) empha-
sized that blends of highly adapted cultivars will seldom
be more stable than the best or better component. Re-
ports of mixtures of barley (Hordeum vulgare L.) with
yields higher than the highest yielding component were
very few (Jedel et al., 1998). In most previous studies,
yields of mixtures have been neutral, in other words
about the same as, or slightly higher than, that of the
weighted means of their components (Wolfe, 1985; Joki-
nen, 1991). Hoekstra et al. (1985) compared all two-
component mixtures of seven maize hybrids with their
pure stands under different densities. In both years,
no mixture at a standard agronomic plant population
density yielded significantly more than either of the
two highest yielding hybrids in pure stand. In a 7-yr
experiment in Nepal comparing barley cultivar mixtures
with pure lines, it was found that even during epidemic
years of yellow rust, the disease reduction in the mixture
was not enough to significantly increase the yield of the
mixture above that of the mean yield of the components
(Pradhanang and Sthapit, 1995).
Size differences within plant communities can be
caused by several factors, some of which affect plant
growth rates, while some affect size in other ways. Size
variation in plant communities may develop because of
(i) age differences; (ii) genetic differences; (iii) environ-
mental heterogeneity; (iv) maternal (seed size) effects;
(v) differential effects of herbivores, parasites, or patho-
gens; or (vi) competition. In most cases, size variation
will be the result of interactions among these factors.
Bonan (1988) showed in a model that competing
plants have a higher size inequality than noncompeting
plants. He interpreted these results as indicating that
the greater degree of size inequality among competing
plants than among noncompeting plants reflects, in part,
variation in relative growth rates induced by neighbor-
hood competition. Bonan (1991) further proposed that
increased plant-to-plant variability at higher plant densi-
ties is a direct manifestation of neighborhood compe-
tition.
Stand Uniformity, Yield, and Stress Tolerance
Resource use efficiency of a crop is inversely related
to plant-to-plant variability because the response of
photosynthesis and grain yield plant21 to input of re-
sources (i.e., PPFD, N, water) is curvilinear (i.e., the
law of diminishing returns). For instance, Edmeades
and Daynard (1979) reported a curvilinear relationship
between assimilate flux plant21 at anthesis and grain
yield plant21, with a positive threshold for assimilate
flux at which grain yield was zero. In nonuniform stands
of field crops, bigger or taller plants have a competitive
advantage over the smaller or shorter ones. There is
much evidence that interactions between plants of the
same genotype at commercial production densities are
always undercompensatory; gains from competition fail
to counterbalance losses (Pendleton and Seif, 1962;
Glenn and Daynard, 1974; Hoekstra et al., 1985; Fasoula
and Fasoula, 1997). Pendleton and Seif (1962) studied
the effects of competition between a tall maize hybrid
‘US 13’ and its near-isogenic dwarf version. All combi-
nations of tall and dwarf plants produced lower yields
than tall and dwarf plants grown alone. A row of tall
maize plants bordered by dwarf plants yielded 6% more
than when bordered by tall plants. A row of dwarf plants
bordered by tall plants yielded 30% less than when
bordered by dwarf plants. A study by Glenn and Dayn-
ard (1974) on the effects of plant-to-plant variation on
maize grain yield demonstrated that plant-to-plant vari-
ation per se lowered grain yield. They suggested that
cultural procedures designed to encourage uniform
plant establishment, like uniform seed bed and constant
planting depth, should maximize maize grain yield. Ford
and Hicks (1992) observed a 5% yield reduction in
maize when half of the stand was delayed in sowing by
7 d, and a 12.8% yield reduction when half of the plant
stand was delayed by 14 d of late sowing. The reduction
in yield increased both with an increase in the propor-
tion of the stand that was delayed and with an increase
in plant population density.
Results of our studies indicate that stand uniformity
and stress tolerance are highly associated (Wu, 1998).
In the first experiment, the relationship between stand
uniformity and stress tolerance was investigated in an
older and a more recent hybrid grown at two plant
densities, 3.5 and 11 plants m22. At each density seeds
were sown either all on the same day to produce uniform
stands (control), or on alternative sowing dates of three
to produce nonuniform stands (treatment). At physio-
logical maturity a significant (P , 0.05) density 3 treat-
ment interaction was observed for grain yield, total
aboveground dry matter, and coefficient of variability
(CV) of the individual plant yield and dry matter. The
CV is the most widely used parameter to quantify vari-
ability among individual plants of a crop stand (e.g.,
Edmeades and Daynard, 1979; Barnes, 1982; Fasoula
and Fasoula, 1997). At the lower density, stands of uni-
form sowing and nonuniform sowing did not differ in
leaf area index, total aboveground dry matter, and grain
yield, nor did they differ in the plant-to-plant variability
for these traits. At the higher plant population density,
higher plant-to-plant variability in stands of both hy-
brids was observed. Nonuniform stands yielded less than
uniform stands in both hybrids at the higher plant popu-
lation density. The difference in grain yield between the
newer and the older hybrid was 30% in the uniform
stand and 46% in the nonuniform stand (Fig. 2). Overall,
stand uniformity declined with an increase of interplant
competition, and its impact on grain yield was greater
in the older than in the newer hybrid. In a second experi-
ment, the association between genetic improvement and
stand uniformity among six maize hybrids released from
1959 to 1995 was explored. Hybrids were grown at two
plant population densities (7 and 11 plants m22) and
two soil N levels. Results depicted in Fig. 3 show that
CV was not associated with yield when CV of the crop
stand was ,30%, which indicates that changes in stan-
 TOLLENAAR & WU: YIELD IMPROVEMENT AND GREATER STRESS TOLERANCE IN MAIZE 1601

Fig. 2. (A) Grain yield and (B) coefficient of variation (CV) for individual plant grain yield of an older (Pride 5) and a newer (Pioneer 3902)
maize hybrid grown at two plant population densities and two levels of stand uniformity at Elora, Ontario in 1997. Values are means of four
replications. Capped bars show standard error.

dard deviation were proportional to changes in yield
(i.e., CV 5 standard deviation/mean). In contrast, a
small decline in yield was associated with a steep in-
crease in CV when CV of the crop stand was .30%.
In both studies density-tolerant hybrids exhibited less
plant-to-plant variability than nontolerant ones at plant
densities close to or higher than their optimum densities.
The association between stand uniformity and stress
tolerance in our studies is supported by studies on hybrid
mixtures and maize stands with uneven emergence. In
studies reported by Hoekstra (1981) and Ford and Hicks
(1992), yield reduction of hybrid mixtures and uneven-
emergence stands of maize increased with the increase
of plant population density (Fig. 4). Maize breeders
have obtained significant yield gains when they selected
for reduced barrenness (Troyer and Rosenbrook, 1983;
Troyer, 1994) and a reduced anthesis-to-silking interval
(Bolaños and Edmeades, 1993a, 1993b, 1996; Beck et
1602 CROP SCIENCE, VOL. 39, NOVEMBER–DECEMBER 1999
Fig. 3. The relationship between grain yield per unit area and coeffi-
cient of variation for individual plant grain yield at physiological
maturity for six maize hybrids grown in four environments. Data
points include four hybrids at 11 plants m22 at Cambridge, Ontario
in 1996; six hybrids at 7.0 plants m22 at two soil nitrogen levels at
Cambridge, Ontario in 1996; 11 plants m22 at Cambridge, Ontario
in 1997; and two hybrids at 11 plants m22 at Elora, Ontario in 1997.
The hybrids Pride 5, Warwick 263, Pioneer 3902, and NK 2555
were used in the test at Cambridge in 1996 and 1997; PAG SX111
and Pioneer 3893 were included at Cambridge in 1997. Pride 5
and Pioneer 3902 were used for the test at Elora, and CV values
were taken from four replications for each hybrid.
al. 1996) in stressful environments, which could be con-
sidered specific aspects of the general phenomenon of
plant-to-plant variation. If stand uniformity and grain
yield are highly correlated across a wide range of geno-
types and environments, selection for stand uniformity
may be a means to increase plant tolerance to environ-
mental stresses.
The Potential for Future Genetic Improvement
A retrospective analysis may show factors that have
contributed to the genetic yield improvement, but not
all of those factors may be able to contribute to future
yield improvements. The initial yield improvement of
maize hybrids over open-pollinated varieties was due
to a large extent to heterosis, i.e., an average yield in-
crease of 15% is commonly attributed to heterosis (cf.,
Frey, 1971), but it is not clear whether heterosis has
contributed beyond the initial increase and whether it
will contribute in the future. Duvick (1984) showed that
rate of yield improvement has been similar for inbreds
and hybrids in the U.S. Corn Belt, suggesting that heter-
osis did not contribute to yield improvement after the
introduction of commercial hybrids. The incorporation
of a defensive trait (e.g., pest resistance) is a one-time
contribution to yield and a continuous breeding effort
may be required to maintain yield potential at the same
level in order to stay one step ahead of the pest organism
as it circumvents the plant’s resistance. Several single-
gene traits that have recently been introduced in maize
Fig. 4. Grain yield comparison between uniform and nonuniform
stands. (A) Grain yield of United 106 and Pride 116 in pure stands
and their mixtures of different proportions (7:1, 6:2, 5:3, 4:4, 3:5,
2:6, 1:7) at three plant population densities at Elora in 1978 and
1979. Data from Hoekstra (1981). (B) Grain yield of uniformly
sown stands and non-uniformly sown (50% plants sowed 7 d earlier
and 50% plants sowed 7 d later relative to uniform stands) stands
at four plant population densities averaged over 5 yr in Minnesota.
Data from Ford and Hicks (1992).
through biotechnology fall in this category. Ledent and
Stoy (1988) showed that 70 yr of wheat (Triticum aesti-
vum L.) breeding in Sweden did not seem to have im-
proved grain yield potential as yield among cultivars
representing 70 yr of breeding did not differ when dis-
eases were controlled by chemical means and lodging
was prevented. In a similar vein, improvement in some
physiological traits may constitute a one-time improve-
ment in yield potential (e.g., erect leaf-angle distri-
bution).
In contrast, some of the factors that have shown either
no or little improvement in the past may contribute to
future yield improvement. For instance, early plant
vigor, possibly combined with cold tolerance, could in-
crease PPFD interception during early phases of devel-
opment. Also, maize researchers in China (Zhao and
Li, 1998) have selected for high Pmax and reported that
Pmax during the growing season was 15% greater in se-
lected lines than in the control, which was associated
with an increase in grain yield. In addition, harvest index
could increase if the increase in grain yield would be
 TOLLENAAR & WU: YIELD IMPROVEMENT AND GREATER STRESS TOLERANCE IN MAIZE
proportionally greater than the increase in total dry
matter (i.e., an increase in total aboveground dry matter
does not have to be associated with an increase in leaf
area per plant, as PPFD interception of maize grown
at commercial plant densities is currently close to 95%).
However, the contribution to future yield improvement
of these three factors is likely to be relatively small.
Increased stress tolerance, combined with increased
stand uniformity under stress conditions, will probably
continue to provide the highest potential for yield im-
provement in maize in the next decades. A large propor-
tion of the genetic improvement of maize in the past is
attributable to increased stress tolerance, and improved
stress tolerance can continue to contribute to genetic
gain in the future. Maximum yields obtained under field
conditions represent an indication of genetic yield po-
tential of current maize hybrids (all yields cited herein
are expressed at 0% grain moisture). A farm-scale yield
of 19.6 Mg ha21 has been recorded by a maize producer
in Illinois (Warsaw, 1985) and a yield of 15.5 Mg ha21 has
been reported for large field research plots in Ontario
(Stevenson, 1985). Average yields of maize in the USA
and Ontario during the period those yields were re-
corded were ≈6.0 Mg ha21. Alternatively, yield potential
is indicated by estimates of grain yield obtained under
close to no-stress conditions. Mean yield during a 4-yr
period was 11.5 Mg ha21 of a short-season hybrid grown
in a growth room in which daily accumulated PPFD was
only 50% of average daily accumulated PPFD during
summer months in Ontario (Tollenaar and Migus, 1984).
The radiation use efficiency in this study was 7.6 g of
dry matter per megajoule of incident photosynthetic
active radiation, which is approximately two times
greater than reported values for field-grown maize (Tol-
lenaar and Agulera, 1992). Overall, these results show
that the potential for continued genetic yield improve-
ment through improved stress tolerance is substantial.
REFERENCES
Barnes, A. 1982. Variability and uniformity in vegetable research.
Internal Rep., Natl. Vegetable Res. Stn., Wellesbourne, War-
wick, UK.
Beck, D., J. Bertran, M. Banziger, G.O. Edmeades, J.M. Ribaut, M.
Willcox, S.K. Vasal, and A. Ortega. 1996. Progress in developing
drought and low soil nitrogen tolerance in maize. p. 85–111. In D.
Wilkinson (ed.) Proc. Annu. Corn and Sorghum Res. Conf., 52th.
Chicago, IL. Am. Seed Trade Assn., Washington, DC.
Bolaños, J., and G.O. Edmeades. 1993a. Eight cycles of selection for
drought tolerance in tropical maize. I. Responses in yield, biomass
and radiation utilization. Field Crops Res. 31:253–268.
Bolaños, J., and G.O. Edmeades. 1993b. Eight cycles of selection for
drought tolerance in tropical maize. II. Responses in reproductive
behaviour. Field Crops Res. 31:269–286.
Bolaños, J., and G.O. Edmeades. 1996. The importance of the anthesis-
silking interval in breeding for drought tolerance in tropical maize.
Field Crops Res. 31:233–252.
Bonan, G.B. 1988. The size structure of theoretical plant populations:
Spatial patterns and neighbourhood effects. Ecology 69:1721–1730.
Bonan, G.B. 1991. Density effects on the size structure of annual
plant populations: An indication of neighbourhood competition.
Ann. Bot. 68 (London):341–347.
Crosbie, T.M. 1982. Changes in physiological traits associated with
long-term breeding efforts to improve grain yield of maize. p.
206–233. In H.D. Loden and D. Wilkinson (ed.) Proc. Annu. Corn
1603
and Sorghum Ind. Res. Conf., 37th. Chicago, IL. 5–9 Dec. 1982.
Am. Seed Trade Assn., Washington, DC.
Derieux, M., M. Darrigrand, A. Gallais, Y. Barriere, D. Bloc, and Y.
Montalant. 1987. Estimation du progres genetique realise chez le
mais grain en France entre 1950 et 1985. Agronomie (Paris) 7:1–11.
Duvick, D.N. 1984. Genetic contributions to yield gains of U.S. hybrid
maize, 1930 to 1980. p. 1–47. In W.R. Fehr (ed.) Genetic contribu-
tions to yield gains of five major crop plants. CSSA Spec. Publ. 7.
ASA and CSSA, Madison, WI.
Duvick, D.N. 1992. Genetic contributions to advances in yield of U.S.
maize. Maydica 37:69–79.
Duvick, D.N. 1997. What is yield? p. 332–335. In G.O. Edmeades et
al. (ed.) Developing drought and low N-tolerant maize. CIMMYT,
El Batan, Mexico.
Dwyer, L.M., and M. Tollenaar. 1989. Genetic improvement in photo-
synthetic response of hybrid maize cultivars, 1959 to 1988. Can. J.
Plant Sci. 69:81–91.
Dwyer, L.M., D.W. Stewart, and M. Tollenaar. 1992. Analysis of maize
leaf photosynthesis under drought stress. Can. J. Plant Sci. 72:
477–481.
Earl, H.J., and M. Tollenaar. 1998. Differences in rates of mature leaf
respiration among commercial maize hybrids. Field Crops Res.
59:9–19.
Edmeades, G.O., and T.B. Daynard. 1979. The relationship between
final yield and photosynthesis at flowering in individual maize
plants. Can. J. Plant Sci. 59:585–601.
Fasoula, D.A., and V.A. Fasoula. 1997. Competitive ability and plant
breeding. Plant Breed. Rev. 14:89–138.
Ford, J.H., and D.R. Hicks. 1992. Corn growth and yield in uneven
emerging stands. J. Prod. Agric. 5:185–188.
Frey, K.J. 1971. Improving crop yields through plant breeding. p.
15–58. In J.D. Eastin and R.D. Munson (ed.) Moving off the yield
plateau. ASA Spec. Publ. 20. ASA, CSSA, and SSSA, Madison, WI.
Glenn, F.B., and T.B. Daynard. 1974. Effects of genotype planting
pattern, and planting density on plant-to-plant variability and grain
yield of corn. Can. J. Plant Sci. 54:323–330.
Goodman, M.M. 1990. Genetic and germplasm stocks worth conserv-
ing. J. Hered. 81:11–16.
Hoekstra, G.J. 1981. Performance of pure stands and mixtures of
maize (Zea mays L.) hybrids. M.S. thesis, Univ. of Guelph, ON,
Canada.
Hoekstra, G.J., L.W. Kannenberg, and B.R. Christie. 1985. Grain
yield comparison of pure stands and equal proportion mixtures for
seven hybrids of maize. Can. J. Plant Sci. 65:471–479.
Jedel, P.E., J.H. Helm, and P.A. Burnett. 1998. Yield, quality and
stress tolerance of barley mixtures in central Alberta. Can. J. Plant
Sci. 78:429–436.
Jokinen, K. 1991 Competition and yield advantage in barley–barley
and barley–oats mixtures. J. Agric. Sci. (Helsinki) 63:255–285.
Ledent, J.F., and V. Stoy. 1988. Yield of winter wheat. A comparison
of genotypes from 1910 to 1976. Cereal Res. Commun. 16:151–156.
Mackay, A.D., and S.A. Barber. 1986. Effect of nitrogen on root
growth of two genotypes in the field. Agron. J. 78:699–703.
Marshall, D.R., and A.H.D. Brown. 1973. Stability of performance
of mixtures and multilines. Euphytica 22:405–412.
McCullough, D.E., Ph. Girardin, M. Mihajlovic, A. Aguilera, and M.
Tollenaar. 1994. Influence of N supply on development and dry
matter accumulation of an old and a new maize hybrid. Can. J.
Plant Sci. 74:471–477.
Nissanka, S.P. 1995. The response of an old and a new maize hybrid
to nitrogen, weed and moisture stress. Ph.D. thesis, Univ. of
Guelph, ON, Canada.
Nissanka, S.P., M.A. Dixon, and M. Tollenaar. 1997. Canopy gas
exchange response to moisture stress in old and new maize hybrid.
Crop Sci. 37:172–181.
Pendleton, J.W., and R.D. Seif. 1962. Role of height in corn competi-
tion. Crop Sci. 2:154–156.
Penning de Vries, F.W.T. 1975. The cost of maintenance processes
ion plant cells. Ann. Bot. (London) 39:77–92.
Penning de Vries, F.W.T., A.H.M. Brunsting, and H.H. van Laar.
1974. Products, requirements, and efficiency of biosynthesis: A
quantitative approach. J. Theor. Biol. 45:339–377.
Pradhanang, P.M., and B.R. Sthapit. 1995. Effect of cultivar mixtures
on yellow rust incidence and grain yield of barley in the hills of
Nepal. Crop Prot. 14:331–334.
1604 CROP SCIENCE, VOL. 39, NOVEMBER–DECEMBER 1999
Rajcan, I., and M. Tollenaar. 1999a. Source–sink ratio and leaf senes-
cence in maize. I. Dry matter accumulation and partitioning during
the grain-filling period. Field Crops Res. 60:245–253.
Rajcan, I., and M. Tollenaar. 1999b. Source–sink ratio and leaf senes-
cence in maize. II. Metabolism of nitrogen and soluble carbohy-
drates during the grain-filling period. Field Crops Res. 60:255–265.
Russell, W.A. 1985. Evaluations for plant, ear, and grain traits of maize
cultivars representing different eras of breeding. Maydica 30:85–96.
Stevenson, C.K. 1985 Maximum yield corn research. p. 190–191. In
R.D. Munson (ed.) Physiology, biochemistry, and chemistry associ-
ated with maximum yield corn. Proc. Workshop Foundation for
Agronomic Research and Potash Phosphate Institute. 11–12 Nov.
1985. St. Louis, MO.
Tollenaar, M. 1989. Genetic improvement in grain yield of commercial
maize hybrids grown in Ontario from 1959 to 1988. Crop Sci.
29:1365–1371.
Tollenaar, M. 1991. The physiological basis of the genetic improve-
ment of maize hybrids in Ontario from 1959 to 1988. Crop Sci.
31:119–124.
Tollenaar, M. 1992. Is low plant density a stress in maize? Maydica
37:305–311.
Tollenaar, M., and A. Aguilera. 1992. Radiation use efficiency of an
old and a new maize hybrid. Agron. J. 84:536–541.
Tollenaar, M., A. Aguilera, and S.P. Nissanka. 1997. Grain yield is
reduced more by weed interference in an old than in a new maize
hybrid. Agron. J. 89:239–246.
Tollenaar, M., and L.M. Dwyer. 1998. Physiology of maize. p. 169–204.
In D.L. Smith and C. Hamel (ed.) Crop yield, physiology and
processes. Springer, New York.
Tollenaar, M., L.M. Dwyer, and D.E. McCullough. 1994. Physiological
basis of genetic improvement of corn. p. 183–236. In G.A. Slafer
(ed.) Genetic improvement of field crops. Marcel Dekker, New
York.
Yield of Wheat in the United Kingdom: Recent Advances and Prospects
R. B. Austin*
ABSTRACT
From 1948 to the present, wheat (Triticum aestivum L.) yields in
the UK have increased by an average of 110 kg ha21 each year. This
rate of increase has been at least maintained in recent years. The
greater yields have been associated with the adoption of cultivars of
shorter stature, which are resistant to lodging and reach anthesis ≈ 1 wk
earlier than old cultivars. In the last two decades, most of these
cultivars have carried the rht D1b dwarfing gene. The full yield benefits
from modern cultivars have depended on high rates of N fertilization
and the use of herbicides and effective fungicides. Data from recent
trials with candidate cultivars and F1 hybrids suggest that further
genetic gain in yield will be achieved during the next decade. Improved
crop protection through chemicals may also enable farmers to obtain
greater yields. In the longer term, substantial genetic gain in yield
may be achieved if breeders are able to produce cultivars with faster
growth rates and greater biomass at maturity. One way to achieve
this would be to modify the photosynthetic enzyme rubisco so that
its oxygenase activity is reduced. However, cultivars with potentially
faster growth rates would require even more N fertilizer if their greater
yield potential is to be realized.
I n the United Kingdom, as in many other countries,
yields of wheat have increased steadily during the
R.B. Austin, 15 Wingate Way, Trumpington, Cambridge, CB2 2HD,
UK. Received 28 Dec. 1998. *Corresponding author (r.b.austin@dial.
pipex.com).
Published in Crop Sci. 39:1604–1610 (1999).
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Tollenaar, M., and M. Mihajlovic. 1991. Bromoxynil tolerance during
the seedling phase is associated with genetic yield improvement in
maize. Can. J. Plant Sci. 71:1021–1027.
Tollenaar, M., M. Mihajlovic, and A. Aguilera. 1991. Temperature
response of dry matter accumulation, leaf photosynthesis, and chlo-
rophyll fluorescence in an old and a new maize hybrid during early
development. Can. J. Plant Sci. 71:353–359.
Tollenaar, M., and W. Migus. 1984. Dry matter accumulation of maize
grown hydrioponically under controlled-environment and field
conditions. Can. J. Plant Sci. 64:475–485.
Troyer, A.F. 1994. Breeding early corn. p. 341–396. In A.R. Hallauer
(ed.) Specialty corns. CRC Press, Boca Raton, FL.
Troyer, A.F. 1996. Breeding widely adapted, popular maize hybrids.
Euphytica 92:163–174.
Troyer, A.F., and R.W. Rosenbrook. 1983. Utility of higher plant
densities for corn performance testing. Crop Sci. 23:863–867.
Vyn, T.J., and M. Tollenaar. 1998. Changes in chemical and physical
quality parameters of maize grain during three decades of yield
improvement. Field Crops Res. 59:135–140.
Warsaw, H. 1985. High yield corn production. p. 193–199. In R.D.
Munson (ed.) Physiology, biochemistry, and chemistry associated
with maximum yield corn. Proc. Workshop Foundation for Agro-
nomic Research and Potash Phosphate Institute. 11–12 Nov. 1985.
St. Louis, MO.
Wolfe, M.S. 1985. The current status and prospects of multiline culti-
vars and variety mixtures for disease resistance. Annu. Rev. Phyto-
pathol. 23:251–273.
Wu, J. 1998. On the relationship between plant-to-plant variability
and stress tolerance in maize (Zea mays L.) hybrids from different
breeding eras. M.S. thesis, Univ. of Guelph, ON, Canada.
Zhao, M., and S. Li. 1998. The development of character of plant
type and photosynthetic traits of corn inbred lines in China. Acta
Agric. Sinica 13:1–4.
period since about 1950. The improvement has been
found to be associated with the greater use of N fertil-
izer, the development of shorter and lodging-resistant
cultivars, the use of crop protection chemicals, improved
and more timely cultivation (especially earlier sowing),
and beneficial interactions between the effects of these
factors. During the period of rapid increases in yield,
the question was often asked: can the rate of increase in
yield be sustained? The evidence is in the yield statistics.
Allowing for year to year variation in yield, wheat yield
in the UK has increased by 110 kg ha21 each year since
1950, and the rate of increase shows no sign of decreas-
ing. Nevertheless, many argue that there must be a bio-
logical limit to yield; that weed, disease, and pest control
cannot be better than complete (as it already is on sub-
stantial areas of the UK wheat hectarage); and that
farmers are already applying close to the optimum rates
of fertilizer N, P, and K. My aim here is to examine the
evidence on the basis of recent improvements in wheat
yields, and to speculate about whether substantial fur-
ther improvements could be made and if so, by what
means. The paper is not concerned with wheat quality,
but it is taken that future production must have at least
the same spectrum of quality characteristics as at
present.
Abbreviations: LAI, leaf area index; NIAB, National Institute of
Agricultural Botany; NL, national list; RL, recommended list.