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Europ. J. Agronomy 28 (2008) 597–605

Yield determination in triticale as affected by radiation in

different development phases
Gaspar Estrada-Campuzano a,∗,1 , Daniel J. Miralles a,b , Gustavo A. Slafer c
a Department of Plant Production and IFEVA, Faculty of Agronomy, Avenue San Martin 4453, (C1417 DSE) Buenos Aires, Argentina
b CONICET (National Council of Scientific and Technological Research) Buenos Aires, Argentina
c ICREA (Catalonian Institution for Research and Advanced Studies) and Department of Crop and Forest Sciences,

University of Lleida, Centre UdL-IRTA, Avenue Rovira Roure 191, 25198 Lleida, Spain
Received 24 October 2007; received in revised form 14 January 2008; accepted 18 January 2008

Abstract
The critical period for grain yield determination has not been determined for triticale. We aimed to identify it, determining the relative importance
of both the major yield components and the dry matter acquisition by the spikes at anthesis. A field experiment was carried out with two triticales,
differing in tillering capacity, subjected to shading treatments at five different timings from early tillering to maturity. Results showed that reductions
in grain yield were more significant when shading was imposed during 3 weeks before and 1 week after anthesis. Reductions in grain yield by
shading treatments were associated with lower number of grains per m2 more than with changes in the average grain weight. Reductions in grains
per m2 were due to reductions in the number of fertile florets per spike, affecting grains per spike. The assimilate acquisition by the spikes during the
critical period was a key determinant of floret survival. Grain number per m2 was related with photothermal quotient during 30 days before anthesis
and spike dry weight at anthesis, though the goodness of the prediction compared with wheat, was lowered by poorer grain setting percentage.
© 2008 Elsevier B.V. All rights reserved.

Keywords: X Triticosecale; Grain number; Grain weight; Intercepted radiation; Photothermal quotient; Shading

1. Introduction In wheat, it has been demonstrated that reducing incident

Although yield components in grain crops are generated
throughout the whole crop growing season (Slafer and Rawson,
1994), yield seems to be much more sensitive to changes in avail-
ability of resources in some particular phases than in others.
As number of grains seems more critical for yield determina-
tion than grain weight (e.g. Sadras, 2007; Peltonen-Sainio et
al., 2007; Fischer, 2008) the ontogenic period during which the
number of grains per m2 is determined have been considered
critical for yield (Slafer et al., 1999). Knowing the window time
when grain yield is more strongly determined could be rele-
vant for developing more adequate strategies for improving yield
through either breeding or management (Slafer, 2003).
∗ Corresponding author. Tel.: +54 11 4524 8063; fax: +54 11 4524 8039.
E-mail addresses: gaspar@agro.uba.ar, escg@uaemex.mx
(G. Estrada-Campuzano).
1 Present address: Facultad de Ciencias Agrı́colas, Universidad Autónoma del
estado de México, Campus Universitario El Cerrillo, El Cerrillo Piedras Blancas,
Toluca 50200, Mexico. Tel.: +52 722 2965518; fax: +52 722 2965518.
radiation during ca. 30 days immediately preceding anthesis
caused more significant reductions in grain number than shading
in any other period (main initial results by Fischer, 1985; later
on validated with many different cultivars and background con-
ditions, e.g. Thorne and Wood, 1987; Savin and Slafer, 1991;
Abbate et al., 1997; Demotes-Mainard and Jeuffroy, 2004). In
all these cases, shading during that critical phase reduced spike
dry weight per unit land area at anthesis, resulting in a pro-
portional reduction in the number of grains per m2 . When water
and nutrients are non-limiting for growth crop, number of grains
per m2 can be trustworthily predicted by the photothermal quo-
tient (i.e. the ratio of mean daily intercepted radiation and mean
daily temperature above a base temperature of 4.5 ◦ C, for the 30
days preceding anthesis), when considering a wide range of con-
ditions given by locations, seasons and management practices
(e.g. Magrin et al., 1993), as it had been proposed from shading
experiments (Fischer, 1985; Savin and Slafer, 1991; Abbate et
al., 1995).
Although some previous evidences in triticale (X Triticose-
cale) suggest that a diminished incident radiation before anthesis
reduced grain yield through reductions in the number of grains

1161-0301/$ – see front matter © 2008 Elsevier B.V. All rights reserved.
doi:10.1016/j.eja.2008.01.003
598 G. Estrada-Campuzano et al. / Europ. J. Agronomy 28 (2008) 597–605
per m2 (Cantarero and Badiali, 1998; Aguirre et al., 2006); there
have been no reports of experiments specifically designed to
determine the timing and length of the window of time where
yield is more strongly determined (i.e. the critical period for yield
determination). The aims of this study were (i) to determine the
window of time for grain yield determination in triticale, (ii)
to identify the yield components more sensitive to changes in
photosynthetically active radiation during that period, and (iii)
to evaluate whether the dry matter acquisition by the spikes at
anthesis determines the number of fertile florets and grains in
triticale as it has been observed for wheat.
2. Materials and methods
2.1. General conditions
A field experiment was carried out during the 2005 growing
season in the experimental field of the Department of Plant Pro-
duction, University of Buenos Aires (34◦ 35 S, 58◦ 29 W) in a
silty clay loam soil classified as Vertic Argiudoll according to
the USDA taxonomy.
Seeds of two triticale cultivars (Yagan and Presto) were hand-
sown at a rate of 350 seeds m−2 on 24 June 2005 in plots of
eight rows width, 0.175 m apart, and 1.4 m long, oriented in
N–S direction. The sowing was made using a special proce-
dure maximizing uniformity: seeds (with more than 95% of
germination) were firstly evenly distributed in strips of sticky-
tape of biodegradable paper of the same length of a row. Soil
was refined to minimize interference with seedling emergence
and individual furrows were manually opened where the strips
of tape were placed and covered and lightly compacted them
by hand and wetted the surface to insure prompt imbibition of
seeds.
During the whole crop cycle the plots were irrigated
(250 mm) to complement natural rainfall (407 mm) occurred
during the cycle with the objective of maintaining the soil near
to field capacity. One week before sowing soil samples were
extracted for the top 0.60 m to determine the soil nitrogen and
phosphorous reserves. Nitrogen (determined as nitrate-N) in the
soil was 30 kgN ha−1 at sowing. In two times, one just before
to beginning of tillering and the other at the onset of stem
elongation, nitrogen fertilizer as urea was applied at a rate of
75 kgN ha−1 each time. Phosphorous fertilizer was not applied
as soil P levels at sowing were >30 mg kg−1 .
Weeds were manually removed, and fungicides and insecti-
cides were sprayed to prevent diseases and pests throughout the
crop cycle. At the beginning of stem elongation (growth stage,
GS 32; Zadoks et al., 1974) and flag leaf appearance (GS 41)
nets were installed to prevent lodging.
2.2. Treatments and design
Treatments consisted of the factorial combination of two trit-
icale cultivars and six shading treatments applied throughout
the crop cycle. The two triticale cultivars were chosen from
previous experiments for possessing under high yielding condi-
Fig. 1. Periods during which shadings were imposed. Arrows show the stages
of seedling emergence (EME), beginning of tillering (BT), beginning of stem
elongation (BSE), flag leaf appearance (FLA), anthesis (ANT) and physiological
maturity (PM), averaged across treatments. Subscripts indicate the difference
in days between anthesis and the midpoint of shading period then used as
independent variable in Figs. 3, 5 and 6.
tions different tillering capacity, though similar time (measured
in thermal time) to anthesis (Estrada-Campuzano et al., 2008).
Presto possessed, in an earlier field study under irrigated and fer-
tilized conditions, higher number of spikes per m2 than Yagan
(600 and 450 spikes per m2 , respectively; data yet unpublished).
Shading treatments consisted of a control (C) unshaded which
received the natural incoming radiation of the season and shad-
ing treatments (Sn ) applied during ca. 25 days at different periods
during crop development (Fig. 1). The exception was the last
shading treatment applied during the whole grain-filling period
(ca. 37 days). In terms of phenological stages, the shading treat-
ments were applied from the beginning of to late tillering (S−83 ),
from then to immediately after the beginning of stem elongation
(S−61 ), from then to flag leaf appearance (FLA) (S−36 ), from
FLA to 1 week after anthesis at the beginning effective grain-
filling period (S−8 ), and from anthesis to physiological maturity
(PM) (S19 ) (Fig. 1). The subscripts for the shading treatments
stand for the difference in days between anthesis and the mid-
point of the shading period (Fig. 1). Shading treatments were
achieved by installing black nylon nets 0.2 m above the top of
the canopy, reducing incident radiation (measured by a linear
ceptometer, see below) by 67 ± 0.3% during treatment imposi-
tion. That value of proportion of the incident radiation retained
by the net was obtained as the quotient between the incident
radiation measured below the shadow net and the correspond-
ing value of the incident radiation immediately above the nylon
net.
Treatments (i.e. cultivars × shading) were arranged in a com-
pletely randomized block design with three replicates. ANOVAs
were performed to determine the effect of treatments, and signif-
icant differences among means of treatments were determined
by the least significance difference (LSD) test.
2.3. Measurements
Since the appearance of the third leaf, two plants per plot were
randomly sampled to determine the stage of apex development
to establish the timing of terminal spikelet initiation. Flag leaf
appearance and anthesis (ANT) were registered when 50% of
the plants within each plot exposed flag leaf completely and the
spikes of the main culms had exerted anthers, respectively. To
determine physiological maturity (PM), 20 random spikes per
plot were tagged at anthesis and dry weight of the basal grains
 G. Estrada-Campuzano et al. / Europ. J. Agronomy 28 (2008) 597–605 599

of the four central spikelets was recorded twice weekly from 7
days after anthesis until maximum grain weight was achieved.
The time of PM was defined as when grain growth finished and
determined by fitting a bilinear regression between dry weight
and thermal time from anthesis (see Miralles and Slafer, 1996).
Duration of all phases in this study was expressed in thermal
time, using 0 ◦ C as base temperature.
To determine above-ground dry matter for the whole crop
cycle (AGDM) plant samples of 30 cm in the two central rows
were taken at the PM stage. Dry weight of the samples was
determined after oven-drying samples for three days at 70 ◦ C.
The number of fertile floret and spike dry weight (after 3 days
at 70 ◦ C) were determined on five spikes of main shoots at anthe-
sis. At PM material was separated into main shoots and tillers and
biomass and grain yield and its components determined within
these two categories.
Air temperature and global incident radiation were mea-
sured hourly using a meteorological station (Vantage Pro2,
Davis instruments, CA, USA). Temperature under nylon nets
was recorded during the shading periods utilizing data loggers
(Cavadevices, Buenos Aires, Argentina). In this sense, averag-
ing over all development stages, shading treatments reduced ca.
0.3 ◦ C mean air temperature, respect to the unshaded treatment
(control), determining differences of ca. 30 ◦ Cd (i.e. mostly irrel-
evant in agronomic terms) in duration to anthesis. The red/far-red
ratio was measured (Skye, Skye instruments, Wales, UK) in both
control and shading treatments and as expected (we used neu-
tral nets for this purpose) it was unaltered by the treatments. On
average, the red/far-red ratio measured at the time of anthesis at
the spikes level, at the half height of the plant and at soil level
were 1.06, 0.30 and 0.26 in the controls and 1.06, 0.31 and 0.30
in the shaded plots, respectively.
Photosynthetically active radiation interception (IPAR) was
measured with a linear ceptometer (LI-191 S, LI-COR Inc. Lin-
coln NE, USA) between 12:00 and 14:00 h on clear days. Four
measurements were taken in each replicate. The first measure-
ment was made above the canopy to determine incident PAR
(I0 ). The other three measurements were taken at the soil surface
or following the senescence profile placing the sensor below the
canopy in three positions along the rows (left, centre and right) at
regular intervals while taking the readings to determine transmit-
ted PAR (It ) as indicated by Charles-Edwards and Lawn (1984).
The fraction of PAR intercepted at midday (F) was calculated
as [I0 − It ]/I0 . Daily fraction interception (FD ) was calculated as
(Charles-Edwards and Lawn, 1984):
2F
FD =
1+F
and applied to corresponding daily integrals of PAR to estimate
intercepted PAR (IPAR). Thus, the fraction of PAR intercepted
at midday was corrected using the equation described above to
calculate the daily fraction interception during the whole day.
Daily incident PAR was calculated as the incident total solar
radiation measured with a standard weather station 50 m from
the plots multiplied by 0.48 (Demotes-Mainard and Jeuffroy,
2004). Daily values were summed from emergence to physio-
logical maturity for each plot to obtain accumulated IPARcum
during the crop cycle. Radiation use efficiency (RUE, g MJ−1 )
was calculated for each treatment as the quotient between above-
ground dry matter for the whole cycle (AGDM) and accumulated
IPARcum . The photothermal quotient (MJ m−2 ◦ C−1 ) was calcu-
lated as the daily ratio between IPAR (corrected as was indicated
above) and mean temperature above a base temperature of 4.5 ◦ C
for the 30 days immediately before anthesis (Fischer, 1985). The
temperature values used considered the minor effect of shading.
3. Results
3.1. Cultivar differences
There were no significant differences in time to anthesis
between both cultivars (1682 and 1725 ◦ Cd for Yagan and Presto,
respectively), neither among shading treatments (1740, 1737,
1745 and 1726 ◦ Cd for the treatments S−83 , S−61 , S−36 and
S−8 , respectively) as average of both cultivars. In fact, as was
indicated in the Material and Methods Section, the differences
in temperature below the shading net was near to 0.3 ◦ C, deter-
mining negligible differences respect to the control.
Grain yield in the control treatment (unshaded) was 1137
and 972 g m−2 for Yagan and Presto, respectively. Yagan had its

Table 1
Grain yield and its components; number of grains per unit land area, average grain weight (AGW), spikes per unit land area, grains per spike, spikelets per spike and
maximum number of tillers per unit land area (MNT) for the whole crop and main shoots or tillers of the cultivars Presto and Yagan in control (unshaded) conditions
Cultivars Grain yield Number of grains AGW Number of Number of grains Number of MNT (m−2 )
(g m−2 ) (10−3 m−2 ) (mg grain−1 ) spikes (m−2 ) (spike−1 ) spikelets (spike−1 )

Total
Yagan 1137.0 a 25.8 b 44.2 a 520 b 49.4 a 32.6 a 1035 b
Presto 972.1 b 28.3 a 34.3 b 736 a 38.4 b 31.0 a 1555 a
Main shoot
Yagan 698.6 a 15.1 a 46.4 a 308 a 49.1 a 33.0 a 307 a
Presto 534.4 b 12.9 b 41.3 a 288 a 44.7 a 31.0 a 288 a
Tillers
Yagan 438.4 a 10.6 b 41.3 a 212 b 49.9 a 32.5 a 728 b
Presto 437.7 a 15.4 a 28.5 b 448 a 34.3 b 31.0 a 1267 a

Means with the same letter within each category comparing cultivars are not significantly different for the LSD0.05 test.
600 G. Estrada-Campuzano et al. / Europ. J. Agronomy 28 (2008) 597–605
Fig. 2. Relationships between (a) grain yield or (b) above-ground dry matter
at maturity (AGDM) and intercepted photosynthetically active radiation accu-
mulated during the crop cycle (IPARcum ) for five shading treatments and one
unshaded control in Yagan (open symbols) and Presto (close symbols). Solid
lines stand for the linear regressions. Vertical and horizontal segments represent
LSD0.05 values.
higher yield associated with heavier grains even though it had
slightly less grains per m2 than Presto (Table 1). As expected,
Presto had a larger number of spikes per m2 (and also maximum
number of tillers) than Yagan, though the latter had a much
higher number of grains per spike due to spikelet fertility, as
both had similar number of spikelets per spike (Table 1). The
contribution of the grains of the tiller spikes to the final number of
grains per m2 was much higher in Presto (ca. 55%) than in Yagan
(ca. 40%), being the grains of the tillers mainly responsible for
the differences in average grain weight (Table 1).
3.2. Shading effects on yield and components
Yield variations, due to shading during different crop phases,
were associated with changes in both above-ground dry matter
biomass during the crop cycle (AGDM) (r2 = 0.57, P < 0.001)
and harvest index (r2 = 0.90, P < 0.001). Additionally, varia-
tions in yield were associated with changes in the IPARcum
Fig. 3. Grain yield relative to the unshaded control in each of the shading treat-
ments (see Fig. 1) for the whole crop (a) and that for main shoots (b) or tillers
(c) in Presto (closed symbols) and Yagan (open symbols). Vertical dotted lines
indicate timing of anthesis. Arrows indicate the beginning of stem elongation
(BSE) and flag leaf appearance (FLA). Segments stand for the LSD0.05 value.
Lines averaging the shape of the responses were fitted by eye.
(r2 = 0.69, P < 0.001), as AGDM was more significantly affected
by IPARcum (r2 = 0.90, P < 0.001) than by changes in RUE
(r2 = 0.22, P > 0.05) (Fig. 2).
Grain yield was reduced in all shading treatments. Although
the magnitude of the effect was different depending on the tim-
ing when shading was imposed, there was a noticeable similar
pattern for both cultivars (Fig. 3a). The highest reduction was
observed when shading was imposed from 2 weeks before to 1
week after anthesis (S−8 ; Fig. 3a). Even though shading after
anthesis was much more severe (ca. 50% longer than all other
shading treatments), its impact on yield was remarkably smaller
than shading just before anthesis and quite similar to that of shad-
ing imposed during the first half of the stem elongation phase
(S−36 ). Earlier shading only marginally, or non-significantly,
affected yield (Fig. 3a). It appeared from these results that yield
is more critically determined from the beginning of stem elon-
gation to mid-grain filling than in any other phases (Fig. 3a).
However, as the latter shading treatment was imposed during the
whole grain filling period was impossible to determine exactly
the end of critical period.
In line with their differences in tillering capacity, yield in
Presto (the cultivar with high tillering capacity) was affected
similarly in both main shoots and tillers, while in Yagan (a cul-
tivar with low tillering capacity) yield was only slightly affected
 G. Estrada-Campuzano et al. / Europ. J. Agronomy 28 (2008) 597–605
Fig. 4. Relationships between (a) yield or (b) average grain weight (AGW) and number of grains per m2 for pre-anthesis shading treatments and (c) number of grains
per m2 and (d) AGW for the control (unshaded) and post-anthesis shading treatment in Presto (closed symbols and bars) and Yagan (open symbols and bars). Solid
line fitted by linear regression. Vertical and horizontal segments as in Fig. 2.
in the main shoots and very responsive in the tillers, so that in
the S−8 treatment the yield was negligible in tillers (Fig. 3b and
c).
Considering shading treatments imposed during pre-anthesis,
variations in grain yield were fully explained by changes in the
number of grains per m2 (Fig. 4a), as the effect of treatments
on this component did not result in any sort of compensation
in average grain weight. In fact, AGW was different between
cultivars, having Yagan heavier grains than Presto, but within
each cultivar, this component was not affected by shading treat-
ments (Fig. 4b). Differences in grain weight between cultivars
when unshaded (Table 1) were maintained independently of the
timing of the pre-anthesis shading treatment (Fig. 4b). Shading
from anthesis onwards (S19 ) reduced both grain number per m2
(Fig. 4c) and average grain weight (Fig. 4d) in both cultivars.
Even though the shading was imposed throughout grain fill-
ing, the effect was larger on number of grains per m2 (ca. 20%
for both cultivars; always significant at P < 0.05) than on aver-
age grain weight (14% in Presto and 7% in Yagan; the former
significant at P < 0.1, and non-significant the latter). The small
reduction in average grain weight (7–14%) contrasted notice-
ably with the magnitude of the shading (67% during the whole
post-anthesis period).
Consequently shading treatments reduced the number of
grains per m2 at maturity with a similar trend to that observed in
yield (Fig. 5a), although the magnitude of the effect was skewed
towards pre-anthesis compared to observe in yield, as part of
the S19 effect on yield was through reductions in average grain
601
weight. The S−8 treatment (24 days of duration) caused the high-
est reductions in the number of grains. Similarly to what has been
described for yield, the effects of shading on main shoot grains
in Yagan were smaller than in Presto (Fig. 5b) and the opposite
was evident in tiller grains (Fig. 5c).
The reductions in the total number of grains per m2 were more
associated with changes in grains per spike than in number of
spikes per m2 (Fig. 6). In fact, the number of spikes per m2 was
only affected by the treatment more strongly affecting yield and
number of grains per m2 (S−8 ) and was virtually unresponsive
to any other timing of shading (Fig. 6a). Obviously the number
of spikes per m2 from main shoots was insensitive (Fig. 6b)
as these treatments might hardly affect plant survival, and all
the responses were due to effects on the number of spikes per
m2 from tillers (Fig. 6c). Number of grains per spike was more
consistently affected by shading treatments (Fig. 6d). In line with
what was described for yield and grains per m2 , the number of
grains per spike in main shoots was only slightly affected in
Yagan by shading and only in the S−8 treatment, while it was
reduced more significantly in Presto (Fig. 6e). Conversely, grains
per spike from tillers were more strongly reduced in Yagan than
in Presto (Fig. 6f).
3.3. Photothermal quotient during the critical period and
spike growth as determinants of grains per m2
Following what is expected from theory and empirical evi-
dences in wheat, grain number per m2 was related to the
602 G. Estrada-Campuzano et al. / Europ. J. Agronomy 28 (2008) 597–605
Fig. 5. Number of grains per m2 , relative to the unshaded control, in each of
the shading treatments (see Fig. 1) for the whole crop (a) and that for main
shoots (b) or tillers (c) in Presto (closed symbols) and Yagan (open symbols).
Vertical dotted lines indicate timing of anthesis. Lines averaging the shape of
the responses were fitted by eye. In the case of panel “a”, the shape was copied
from than reported by Fischer (1985) for bread wheat. Segments as in Fig. 3.
photothermal quotient during the critical period for grain num-
ber determination (Fig. 7). Additionally, the number of grains
per unit area was linearly and positively associated (r2 = 0.32,
P < 0.05) with the spike dry matter per unit area at anthesis. The
partitioning of AGDM to the spike at anthesis (0.16 g g−1 ), cal-
culated as the slope of the relationship between spike dry weight
and biomass at that stage, was maintained constant (r2 = 0.90,
P < 0.001) independently of cultivar or shading treatment (data
not shown).
The relationship between the number of grains per m2 and
spike dry weight at anthesis, although significant, was poorer
than expected from literature reports on wheat. The number of
grains depends upon the number of fertile florets at anthesis
and the efficiency of these florets for setting grains afterwards.
Although a number of fertile florets were only measured in
main shoot spikes it seemed clear that part of the lack of
a better adjustment between the number of grains and spike
dry weight was due to a relatively large abortion (fertile flo-
rets not setting grains). In fact, the relationship between the
number of fertile florets was quite closely associated with the
spike dry weight at anthesis (Fig. 8a). For instance, spike dry
weight to anthesis was reduced by the S−8 treatment 33 and
22% for Presto and Yagan, respectively, and the fertile flo-
rets to anthesis were affected almost in the same magnitude
(36 and 24% in Presto and Yagan, respectively; Fig. 8a). The
relationship of the number of grains per spike with the spike
dry weight loose part of the matching (Fig. 8a) due to an
increased abortion when the number of fertile florets increased
(Fig. 8b).
4. Discussion
The results of the present work demonstrated that triticale
yield was particularly sensitive to reductions in radiation levels
during few weeks immediately before to few days after anthesis
(i.e. from beginning of stem elongation to mid-grain filling).
This is in agreement with reports for other cereals such as wheat
(Fischer, 1985; Savin and Slafer, 1991; Zhu et al., 2004), maize
(Kiniry and Ritchie, 1985; Otegui and Andrade, 2000) and barley
(Arisnabarreta and Miralles, yet unpublished). To the best of our
knowledge shading experiments in triticale were only conducted
with a single pre-anthesis treatment (virtually during the whole
stem elongation phase; Cantarero and Badiali, 1998; Aguirre et
al., 2006). Although, these experiments showed that yield was
sensitive to shading during pre-anthesis, they could not define
whether this sensitivity was of different magnitude to that of
earlier phases.
As expected, reductions in grain yield by pre-anthesis shading
treatments were explained by reductions in number of grains per
m2 without compensations in average grain weight. Both aspects
are in line with the case most commonly reported in wheat (and
barley) where grain number per m2 is the main yield component
explaining changes in yield due to genetic and management fac-
tors (e.g. Slafer et al., 2005; Sadras, 2007; Peltonen-Sainio et
al., 2007; and references quoted therein). Grain weight in cere-
als seems to be strongly sink-limited (e.g. Borrás et al., 2004;
Bingham et al., 2007a) and therefore no clear compensations
are expected from reductions in the number of grains growing
after anthesis. Shading during post-anthesis also affected grain
number per m2 more strongly than average grain weight. The
effect of shading immediately after anthesis on grain number due
to its effects in grain setting has been reported in wheat (Savin
and Slafer, 1991), and lack of major effects of post-anthesis
shading on average grain weight has been reported not only in
wheat (Fischer, 1985; Savin and Slafer, 1991) but also in triti-
cale (Cantarero and Badiali, 1998; Aguirre et al., 2006). In fact,
the minor effect on grain weight found in this study might well
be due to a small reduction in potential grain size (Bingham et
al., 2007b) than a reduction in availability of assimilates needed
to match grain growth requirements during the effective grain
filling period, as was reported in wheat (Slafer and Savin, 1994)
and barley (Voltas et al., 1997). This lack of responsiveness on
average grain weight to intense shading during grain filling rein-
forces the conclusion that grain growth in triticale would be
strongly sink-limited, like it is the most common case in wheat
and barley.
Naturally due to its strong relationship with yield, the number
of grains per m2 exhibited a critical window time similar to that
described for yield, though slightly skewed towards pre-anthesis.
Thereby, it can be concluded that the yield in triticale is more
sensitive to reductions in incident radiation during the period
that comprised between flag leaf appearance and 10 days after
anthesis, very similar to that observed in wheat (Fischer, 1985).
 G. Estrada-Campuzano et al. / Europ. J. Agronomy 28 (2008) 597–605 603

Fig. 6. Spikes per m2 (a–c) and grains per spike (d–f) relative to the unshaded control in each of the shading treatments (see Fig. 1) for the whole crop (a and b) and
for main shoots (c and d) or tillers (e and f) in Presto (closed symbols) and Yagan (open symbols). Vertical dotted lines indicate timing of anthesis. Segments as in
Fig. 3. Lines averaging the shape of the responses were fitted by eye.

In this study, the number of grains per m2 was related to

Fig. 7. Relationship between number of grains per m2 and the photothermal
quotient during the 30 days immediately before to anthesis. Solid line fitted by
linear regression. Vertical and horizontal segments as in Fig. 2.
photothermal quotient, an integrative variable of the growth
and developmental process that are occurring during the spike
growth period (i.e. during ca. 30 days before to anthesis) and
with the spike dry weight at anthesis in line with previous evi-
dences reported in other cereals as wheat (Fischer, 1985; Savin
and Slafer, 1991; Abbate et al., 1995). However, for triticale
(at least for the data set used in this study), these relation-
ships were weaker, likely due to the high percentage of floret
abortion determining an increased reduction in grain setting as
the number of fertile florets at anthesis was increased, coun-
terbalancing the advantages of producing more fertile florets at
anthesis, similarly to that observed in wheat by González et al.
(2005).
We selected two cultivars with different structure of genera-
tion of grain number per m2 where one of them (Yagan) had a
stronger dependence on the grains produced in the main shoot
with low dependence on tillers productivity, while the other cul-
tivar (Presto) had a stronger contribution from tillers with less
dependence on main shoot productivity. In line with these char-
acteristics, they strongly differed in terms of the sub-components
that were most affected by treatments. However, the fact that this
differential behavior in sub-components did not result in any
noticeable differential response of the number of grains per m2 ,
604 G. Estrada-Campuzano et al. / Europ. J. Agronomy 28 (2008) 597–605
Fig. 8. Relationships between (a) the number of fertile florets per main shoot spike and spike dry weight at anthesis and (b) the number of grains and the number of
fertile florets per main shoot spike in two triticale cultivars for different pre-anthesis shading treatments. Solid lines fitted by linear regression. Vertical and horizontal
segments as in Fig. 2. Dashed line in panel b corresponds to the 1:1 ratio, y = x. Please note that the scales do not start in zero.
support that the window time for the number of grains (and yield)
determination is unaltered by the structure of sub-components
determining grain number.
Acknowledgements
We gratefully acknowledge the excellent technical assistance
during the field experiment of I. Alzueta and J. Martinez. G.
Estrada-Campuzano held an overseas PhD Scholarship from
Autonomous University of the State of Mexico (UAEMex) and
Mexican Government (PROMEP). This study was partially sup-
ported by UBACyT (G 067) and FONCYT (PICT 08-13935)
competitive grants.
References
Abbate, P.E., Andrade, F.E., Culot, J.P., 1995. The effects of radiation and
nitrogen on number of grains in wheat. J. Agric. Sci. Camb. 124, 351–360.
Abbate, P.E., Andrade, F.H., Culot, J.P., Bindraban, P.S., 1997. Grain yield in
wheat: effects of radiation during spike growth period. Field Crops Res. 54,
245–257.
Aguirre, A., Rubiolo, O.J., Ribotta, P.D., Lujan, J.S., Perez, G.T., Leon, A.E.,
2006. Effects of incident radiation and nitrogen availability on the quality
parameters of triticale grains in Argentina. Exp. Agric. 42, 311–322.
Bingham, I.J., Blake, J., Foulkes, M.J., Spink, J., 2007a. Is barley yield in the UK
sink limited? I. Post-anthesis radiation interception, radiation use efficiency
and source-sink balance. Field Crops Res. 101, 198–211.
Bingham, I.J., Blake, J., Foulkes, M.J., Spink, J., 2007b. Is barley yield in the
UK sink limited? II. Factors affecting potential grain size. Field Crops Res.
101, 212–220.
Borrás, L., Slafer, G.A., Otegui, M.E., 2004. Seed dry weight response to source-
sink manipulations in wheat, maize and soybean: a quantitative reappraisal.
Field Crops Res. 86, 131–146.
Cantarero, M.G., Badiali, O.J.J., 1998. Modificación de la cantidad de radiación
en pre y post antesis en triticale: Rendimiento en grano y componentes.
In: IV Congreso Nacional de trigo y II Simposio Nacional de cereales de
siembra otoño—invernal, Mar del Plata, Argentina.
Charles-Edwards, D.A., Lawn, R.J., 1984. Light interception by grain legume
row crops. Plant Cell Environ. 7, 241–251.
Demotes-Mainard, S., Jeuffroy, M.H., 2004. Effects of nitrogen and radiation
on dry matter and nitrogen accumulation in the spike of winter wheat. Field
Crops Res. 87, 221–233.
Estrada-Campuzano, G., Miralles, D.J., Slafer, G.A., 2008. Genotypic variability
and response to water stress of pre and post anthesis phases in triticale. Eur.
J. Agron. 28, 171–177.
Fischer, R.A., 1985. Number of kernels in wheat crops and the influ-
ence of solar radiation and temperature. J. Agric. Sci. Camb. 105, 447–
461.
Fischer, R.A., 2008. The importance of grain or kernel number in wheat: a reply
to Sinclair and Jamieson. Field Crops Res. 105, 15–21.
González, F.G., Slafer, G.A., Miralles, D.J., 2005. Photoperiod during stem elon-
gation in wheat: is its impact on fertile floret and grain number determination
similar to that of radiation? Func. Plant Biol. 32, 181–188.
Kiniry, J.R., Ritchie, J.T., 1985. Shade-sensitive interval of kernel number of
maize. Agron. J. 77, 711–715.
Magrin, G.O., Hall, A.J., Baldy, C., Grondona, M.O., 1993. Spatial and interan-
nual variations in the photothermal quotient: implications for the potential
kernel number of wheat crops in Argentina. Agric. Forest Meteorol. 67, 29–
41.
Miralles, D.J., Slafer, G.A., 1996. Grain weight reductions in wheat associ-
ated with semidwarfism: an analysis of grain weight at different positions
within the spike of near-isogenic lines. J. Agron. Crop Sci. 177, 9–
16.
Otegui, M.E., Andrade, F.E., 2000. New relationships between light intercep-
tion, ear growth, and kernel set in maize. In: Agronomy, C.S.S.o.A.a.A.S.o.
Physiology and modeling Kernel Set in Maize. CSSA, Madison, pp. 89–
102.
Peltonen-Sainio, P., Kangas, A., Salo, Y., Jauhiainen, L., 2007. Grain number
dominates grain weight in temperate cereal yield determination: evidence
based on 30 years of multi-location trials. Field Crops Res. 100, 179–
188.
Sadras, V.O., 2007. Evolutionary aspects of the trade-off between seed size and
number in crops. Field Crops Res. 100, 125–138.
Savin, R., Slafer, G.A., 1991. Shading effects on the yield of an Argentinian
wheat cultivar. J. Agric. Sci. Camb. 116, 1–7.
Slafer, G.A., 2003. Genetic basis of yield as viewed from a crop physiologist’s
perspective. Ann. Appl. Biol. 142, 117–128.
Slafer, G.A., Araus, J.L., Richards, R.A., 1999. Physiological traits that increase
the yield potential of wheat. In: Satorre, E.H., Slafe, G.A. (Eds.), Wheat:
Ecology and Physiology of Yield Determination. Food Product Press, New
York, pp. 379–416.
Slafer, G.A., Araus, J.L., Royo, C., Garcia del Moral, L.F., 2005.
Promising eco-physiological traits for genetic improvement of cereal
yields in Mediterranean environments. Ann. Appl. Biol. 146, 61–
70.
Slafer, G.A., Rawson, H.M., 1994. Sensitivity of wheat phasic development to
major environmental factors: a re-examination of some assumptions made
by physiologist and modelers. Aust. J. Plant Physiol. 21, 393–426.
Slafer, G.A., Savin, R., 1994. Source-sink relationships and grain mass at
different positions within the spike in wheat. Field Crops Res. 37, 39–
49.
Thorne, G.N., Wood, D.W., 1987. Effects of radiation and temperature on
tiller survival, grain number and grain yield in winter wheat. Ann. Bot.
59, 413–426.
 G. Estrada-Campuzano et al. / Europ. J. Agronomy 28 (2008) 597–605 605

Voltas, J., Romagosa, I., Araus, J.L., 1997. Grain size and nitrogen accumulation
in sink-reduced barley under Mediterranean conditions. Field Crops Res. 52,
117–126.
Zadoks, J.C., Chang, T.T., Konzak, C.F., 1974. A decimal code for the growth
stage of cereals. Weed Res. 14, 415–421.
Zhu, G.X., Midmore, D.J., Radford, B.J., Yule, D.F., 2004. Effect of timing
of defoliation on wheat (Triticum aestivum) in central Queensland. I. Crop
response and yield. Field Crops Res. 88, 211–226.