Field Crops Research 241 (2019) 107575

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Field Crops Research

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Yield determination and the critical period of faba bean (Vicia faba L.) T
a,⁎ b b c
Lachlan Lake , Diego E. Godoy-Kutchartt , Daniel F. Calderini , Andrew Verrell ,
Victor O. Sadrasa,d
a
South Australian Research and Development Institute, Waite Campus, Australia
b
Institute of Plant Production and Protection, Universidad Austral de Chile, Valdivia, Chile
c
Department of Primary Industries, Tamworth Agricultural Institute, Australia
d
School of Agriculture, Food and Wine, The University of Adelaide, Australia

ARTICLE INFO ABSTRACT

Keywords: Faba bean is a cool-season, indeterminate grain legume grown in Mediterranean and temperate environments
Yield determination where yield loss associated with abiotic and biotic stress is common. Yield responses depend on the timing of
Phenology stress relative to the species-specific vulnerable stages in the crop lifecycle. To determine the critical period for
Seed number yield determination in faba bean, we used sequential 14-d shading periods in locally-adapted varieties grown in
Seed size
five environments with yield of unshaded controls from 2.3 to 6.8 t ha−1.
Stress
Yield of shaded treatments diverged from the controls around 450 °Cd before flowering, reached the largest
difference around 100 °Cd after flowering, and converged with controls towards 700–800 °Cd after flowering; the
most critical stage aligned with pod emergence. Seed number accounted for most of the variation in yield
response to shading. Shading increased seed size when stress severely reduced seed number. Pod number ac-
counted for most of the variation in seed number for shading before flowering, and both pod number and seeds
per pod contributed to the reduction in seed number in crops shaded after flowering. The defined critical period
is a useful reference for breeding and agronomic solutions to improve yield under stress.

1. Introduction
Faba bean (Vicia faba L.) is a high-protein, cool-season, in-
determinate grain legume with benefits associated with symbiotic ni-
trogen fixation and improved rotational and weed control options
(Ghanem et al., 2015; Catt and Paull, 2017; Redden et al., 2018). It is
grown predominantly in rainfed cropping systems where a range of
biotic and abiotic stresses lead to low and unreliable yield constraining
production, profitability and adoption (Mwanamwenge et al., 1999;
Siddique et al., 2001; Bhandari et al., 2016; Bodner et al., 2018).
Grain yield is associated with crop growth rate in species-specific
critical developmental periods (Andrade et al., 2005; Sadras et al.,
2013; Lake and Sadras, 2016). Critical periods have been determined in
cereals including wheat, barley, oat, rice, maize, triticale and sorghum
(Fischer, 1985; Kiniry and Ritchie, 1985; Savin and Slafer, 1991;
Arisnabarreta and Miralles, 2008; Estrada-Campuzano et al., 2008;
Oosterom and Hammer, 2008; Cerrudo et al., 2013; Mahadevan et al.,
2016; Ali et al., 2019), sunflower (Cantagallo et al., 1997), canola
(Kirkegaard et al., 2018), quinoa (Bertero and Ruiz, 2008) and grain
legumes including lupin, field pea, chickpea and soybean (Board and
Tan, 1995; Jiang and Egli, 1995; Kantolic and Slafer, 2001; Guilioni
et al., 2003; Sandaña et al., 2009; Sandaña and Calderini, 2012; Lake
and Sadras, 2014).
Whereas critical periods are species specific, there is a pattern as-
sociated with growth habit. In the determinate cereals the critical
period occurs in the stages leading into flowering, while in the studied
indeterminate grain legumes and canola the critical period is displaced
towards pod set (Andrade et al., 2005; Sandaña and Calderini, 2012;
Sadras and Dreccer, 2015; Kirkegaard et al., 2018).
Diverse stresses including drought and nutrient deficiency primarily
reduce the rate of canopy expansion and, with more severe stress, ra-
diation use efficiency. Sequential periods of shade over the duration of
the growth cycle capture these primary effects, and have been com-
monly used to reliably determine the critical period in different crops
(Arisnabarreta and Miralles, 2008; Sandaña and Calderini, 2012; Lake
and Sadras, 2014; Mahadevan et al., 2016; Kirkegaard et al., 2018).
Studies on water stress suggest that the critical period of faba bean may
be around podding (Mwanamwenge et al., 1999; Bodner et al., 2018;
Zeleke and Nendel, 2019), but a targeted experiment is yet to be un-
dertaken. The aim of this study was to quantify the critical period for
yield determination in faba bean and the associated contributions of the
underlying yield components.

⁎
Corresponding author.
E-mail address: lachlan.lake@sa.gov.au (L. Lake).

https://doi.org/10.1016/j.fcr.2019.107575
Received 7 May 2019; Received in revised form 14 July 2019; Accepted 14 July 2019
Available online 18 July 2019
0378-4290/ © 2019 Elsevier B.V. All rights reserved.
L. Lake, et al. Field Crops Research 241 (2019) 107575

Fig. 1. Cumulative rainfall and evaporation, and daily radiation and temperature during the growing season in five environments. The phenological scale is for the
unshaded controls.

2. Methods 2.2. Treatments and experimental design

2.1. Environments and varieties
Crops were grown using local practices for establishment and con-
trol of diseases, pests and weeds in five environments resulting from the
combination of locations and seasons. Locations were Roseworthy in
South Australia (34°52′S, 138°69′E), Tamworth (34°33′S, 138°49′E) and
Breeza (31°27′S, 150°46′E) in New South Wales, and Valdivia in
southern Chile (39°47′S, 73° 14′W). Crops were sown on the 20th of
June 2017 and 6th of June 2018 at Roseworthy, 3rd of May 2017 at
Tamworth, 19th of June 2018 at Breeza, and 14th August 2018 in
Valdivia. Sowing density ranged from 30 seeds m−2 (Australia) to 45
seeds m−2 (Valdivia). Daily rainfall, reference evapotranspiration,
maximum and minimum temperature and radiation were obtained from
nearby weather stations (Fig. 1).
Roseworthy crops were sown after barley (2017) and wheat (2018)
in a Calcic Luvisol, following wheat on a red chromosol at Tamworth,
following wheat on a black vertosol at Breeza, and a Duric Hapludand
volcanic soil in Valdivia. Seed was inoculated only where background
soil Rhizobium were not sufficient for nodulation.
Locally-adapted varieties of varying seed size and phenology were
PBA Rana and AFO 3001-1 in Roseworthy; PBA Nasma and PBA Warda
in Tamworth and Breeza; Histal and Portuguesa in Valdivia (Table 1).
Table 1
Duration of the period between sowing and flowering of locally-adapted faba
bean varieties in five environments. Data from unshaded controls.
Environment Variety Seed Duration sowing Duration
size to flowering flowering to
(mg) maturity
The trials were arranged in a split-plot design with three replicates,
where varieties were allocated to main plots and 14-d sequential
shading treatments, including unshaded controls, to randomised sub-
plots. Shades were constructed from black shade cloth that intercepted
around 90% (Australia) and 75% (Chile) of photosynthetically active
radiation PAR (measured with a ceptometer), and were maintained at a
minimum of 10 cm from the top of the canopy at all times. The shade
cloth was attached to a frame using wire and metal so that plants were
shaded from the top and three sides, with the southern side left open to
allow for air flow, regular variation in temperature and relative hu-
midity and access to pollinators (Lake and Sadras, 2014; Kirkegaard
et al., 2018). To ensure that temperature was not a confounding factor,
we recorded temperature inside and outside the shade structures at
Roseworthy using HOBO MX100 data loggers. Temperature was logged
every 20 min and we found that daily differences were less than the
quoted accuracy of the loggers (< 0.5 °C).
The number of shading treatments varied between years and en-
vironments due to variation in sowing date and duration of the growing
season; the latest treatment included the brown-pod stage corre-
sponding with the final stage of seed abortion just before physiological
maturity (Ney and Turc, 1993; Munier-Jolain et al., 1998).
At Roseworthy, main plot size was 65 m2, comprised of 6 rows
(spaced 24 cm) of 45-m length; subplots were 1.2 m2 comprised of 5
rows of 1-m length. At Tamworth, main plot size was 91 m2, comprised
of 5 rows (spaced 38 cm) of 48-m length; subplots were 1.14 m2 com-
prised of 3 rows of 1-m length. At Breeza, main plot size was 72 m2,
comprised of 3 rows (spaced 50 cm) of 48-m length on 2 m wide raised
irrigation beds; subplots were 1.5 m2 comprised of 3 rows of 1-m
length. In Valdivia, individual plots consisted on 5 rows, 1.6 m long and
0.15 m apart.

(d) (°Cd) (d) (°Cd)

2.3. Traits
Roseworthy 2017 PBA Rana 69 81 970 54 831
AFO 3001-1 56 82 988 54 834
We monitored phenology weekly to determine when fifty percent of
Roseworthy 2018 PBA Rana 62 79 898 56 774
AFO 3001-1 54 77 873 58 794 the plants within the plot had reached: flowering (open petals), pod
Tamworth 2017 PBA Warda 46 103 1134 70 1050 emergence (pods < 10 mm visible), end of flowering and maturity
PBA Nasma 57 103 1134 70 1050 (Knott, 1990; Lake and Sadras, 2014). Phenological stages are ex-
Breeza 2018 PBA Warda 39 83 956 65 1272 pressed on a thermal time scale with base temperature of 0 °C (Catt and
PBA Nasma 45 83 956 65 1272
Paull, 2017).
Valdivia 2018 Histal 140 84 803 50 704
Portuguesa 186 79 748 55 758 Yield and yield components were measured at maturity in shoot
samples from 3 × 0.5 m cuts in central rows (2 in Breeza 2018); border

2
L. Lake, et al.
rows were excluded (Rebetzke et al., 2014). Samples were dried for
72 h at 70 °C or until constant weight. Yield components included shoot
biomass (t ha−1), harvest index (seed yield/shoot biomass), seed
number (m−2), individual seed size (mg), pod number (pods m−2), and
seeds per pod.
2.4. Data analyses
The effects of timing of shading, variety and their interaction were
tested using analysis of variance separately for each environment as
there was an unequal number of shading treatments among environ-
ments. Fisher’s PSLD test was used to determine differences between
shading treatment and unshaded controls.
Yield and yield components in shading treatments were normalised
as a fraction of the control, and the trajectory of normalised traits was
plotted against the phenology of controls (thermal-time scale centred at
flowering). Normalising the treatment values against the controls shows
the relative change of yield and its components across environment
thus allowing environments with different yield potential to be more
readily compared; this robust approach has been used in wheat
(Fischer, 1985), barley (Arisnabarreta and Miralles, 2008), maize
(Kiniry and Ritchie, 1985), sunflower (Cantagallo et al., 1997), field pea
and lupin (Sandaña and Calderini, 2012), canola (Kirkegaard et al.,
2018),and chickpea (Lake and Sadras, 2014). Curves were fitted using
polynomial regression where shading effect was significant and to the
average ratio where non-significant (Arisnabarreta and Miralles, 2008;
Estrada-Campuzano et al., 2008; Sandaña and Calderini, 2012; Lake
and Sadras, 2014). We define the critical period as the period where
yield of treatments diverges significantly from the controls.
3. Results
3.1. Environmental conditions and phenology
Seasonal rainfall exceeded seasonal reference evapotranspiration in
Valdivia, while evaporation exceeded rainfall in the Australian loca-
tions (Fig. 1). Rainfall matched reference evapotranspiration until
flowering at Roseworthy 2017, and crops grew under shortage of
rainfall during the whole season at Breeza 2018. Crops at Tamworth
2017 experienced several frost events (Fig. 1), and high temperature
over 30 °C were common in the reproductive phase for all sites except
Valdivia (Fig. 1), where average temperature is typically 15 °C during
the reproductive phase.
Time from sowing to flowering ranged from 77 to 103 d, and 748 to
1134 °Cd (Table 1). Time from sowing to maturity ranged from 136 to
182 d, and 1662 to 2324 °Cd.
3.2. Seed yield and components
Yield of unshaded controls ranged from 2.3 t ha-1 in Tamworth
2017 to 6.8 t ha−1 in Valdivia, in line with the variation in rainfall,
reference evapotranspiration and average temperature (Fig. 1).
In all environments shade significantly reduced yield (P < 0.05).
Variety did not affect yield in any of the environments nor was there
any interaction between shade and variety.
Owing to both intra-seasonal variation in PAR in a given environ-
ment and variation between environments (Fig. 1), PAR underneath the
shades differed between timing of treatments and between environ-
ments. The relationship between yield reduction and PAR within the
shade period, however, was non-significant within and across en-
vironments (P > 0.05). Likewise, there was no relationship between
3
Field Crops Research 241 (2019) 107575
yield reduction and temperature or rainfall within the shade period
(P > 0.05).
Table 2 presents the matrix of correlations between yield and
components. Yield was strongly correlated with biomass, and more
weakly associated with harvest index. Biomass and harvest index were
unrelated, except at Breeza 2018 where they correlated positively. Yield
correlated with seed number across the environments and varieties, and
was unrelated or weakly related with seed size. Seed number was more
closely associated with pod number than with seeds per pod, except in
Valdivia where seeds per pod was more relevant. There was a trade-off
between seed number and seed size in all environments except Breeza
2018, and between seed size and seeds per pod in three out of five
environments.
3.3. Critical period
Fig. 2a shows yield response to timing of shading across varieties
and environments. Yield of shaded treatments diverged from the con-
trols around 450 °Cd before flowering, reached the largest difference
around 100 °Cd after flowering, and converged with controls towards
700–800 °Cd after flowering. The critical period for yield determination
was therefore 1250 °Cd-long centred 100 °Cd after flowering, corre-
sponding to the time of pod emergence.
Seed number mediated the yield response to shading (Fig. 2b).
Shading centred at 53 °Cd after flowering reduced seed size at Rose-
worthy 2017. In all other cases, shading did not affect or increased seed
size (Fig. 2c).
The reduced seed number was mainly accounted for by decreased
pod number with shading before flowering, and by a combination of
reduced pod number and fewer seeds per pod with shading after
flowering (Fig. 3a and b).
4. Discussion
Shading has been widely used to determine the effect of timing of
stress on yield and its components (Fischer, 1985; Kiniry and Ritchie,
1985; Egli and Yu, 1991; Cantagallo et al., 1997; Arisnabarreta and
Miralles, 2008; Sandaña and Calderini, 2012; Lake and Sadras, 2014;
Mahadevan et al., 2016; Kirkegaard et al., 2018). This approach is fa-
voured because it is highly repeatable and captures the effects of
multiple stresses that are mediated by changes in crop growth rate.
Critical periods determined with this method can be extrapolated to
other stress such as water and nutrient deficit, as yield is primarily
determined by seed number and seed number correlates with crop
growth rate in the critical period irrespective of the driver (Egli and Yu,
1991; Tollenaar et al., 1992; Guilioni et al., 2003; Andrade et al., 2005;
Arisnabarreta and Miralles, 2008; Bertero and Ruiz, 2008; Sadras and
Lawson, 2011; Sandaña and Calderini, 2012). Examples in wheat and
maize reinforce this proposition as the relationship between growth in
the critical period and seed number is maintained irrespective of the
sources of variation in growth including lodging, nitrogen, radiation,
shading or water (Fischer, 1985; Savin and Slafer, 1991; Fischer, 1993;
Prystupa et al., 2004; Andrade et al., 2005; Acreche and Slafer, 2011).
A consideration to using shade as a stress is the daily variation in
solar radiation and the seasonal trends affecting the relative reduction
in radiation compared to controls. In this research we overcome this by
using high radiation intercepting cloth, and numerous environments
with the resulting critical periods showing conformity. The magnitude
of shade-induced yield reduction was larger in Australian environments
(90% shade) than in Chile (75% shade) but the trends in reduction were
consistent irrespective of magnitude. Indeed, the method focuses on the
L. Lake, et al. Field Crops Research 241 (2019) 107575

Table 2
Correlation matrix of yield and its components. Correlations are based on averages of two varieties, split by environment.
Seed number Seed size Biomass Harvest Index Seeds per pod Pod number

Roseworthy 2017
Yield 0.90*** 0.11 0.76*** 0.50*** 0.30* 0.87***
Seed number −0.30* 0.51*** 0.69*** 0.45** 0.92***
Seed size 0.46** −0.44** −0.42** −0.19
Biomass −0.15 −0.10 0.63***
Harvest Index 0.55*** 0.50***
Seeds per pod 0.08

Roseworthy 2018
Yield 0.81*** 0.28* 0.88*** 0.54*** 0.07 0.93***
Seed number −0.31* 0.63*** 0.57*** 0.53*** 0.86***
Seed size 0.41** −0.08 −0.76*** 0.09
Biomass 0.09 −0.15 0.83***
Harvest Index 0.38** 0.47***
Seeds per pod 0.05

Tamworth 2017
Yield 0.85*** −0.02 0.91*** 0.59*** 0.19 0.82***
Seed number −0.52*** 0.74*** 0.51*** 0.26* 0.97***
Seed size 0.03 −0.07 −0.18 −0.50***
Biomass 0.21 0.16 0.73***
Harvest Index 0.12 0.50***
Seeds per pod 0.01

Breeza 2018
Yield 0.95*** 0.32* 0.97*** 0.80*** 0.41** 0.93***
Seed number 0.07 0.94*** 0.72*** 0.42** 0.99***
Seed size 0.30* 0.30* 0.17 0.02
Biomass 0.63*** 0.39** 0.92***
Harvest Index 0.39** 0.69***
Seeds per pod 0.30*

Valdivia 2018
Yield 0.81*** −0.07 0.83*** 0.41* 0.25 0.65***
Seed number −0.60*** 0.55** 0.50* 0.67*** 0.37*
Seed size −0.01 −0.07 −0.70*** 0.04
Biomass −0.17 −0.09 0.84***
Harvest Index 0.57** −0.23
Seeds per pod −0.39*

Significance is ***P < 0.0001, **P < 0.001 and *P < 0.05 according to Fisher’s r to Z test.

timing not the severity of yield reduction.
The critical period for yield determination is an important physio-
logical concept that has been demonstrated in cereals, oilseeds and
some grain legumes. Yield correlates with physiological status during
the critical period, quantified for example as crop growth rate (Andrade
et al., 2005; Egli, 2019); hence the implications for breeding and agr-
onomy.
Critical periods are species specific with generally little variation
within species. However, critical periods cluster with growth habit as
shown in comparisons between determinate cereals which have more
separate vegetative and reproductive phases and indeterminate grain
legumes where the two phases overlap (Sadras and Dreccer, 2015). In
our environments the critical period for faba bean represented between
50 to 75% of the crop cycle (˜1250 °Cd) with some degree of yield loss
occurring for the majority of the growing season. Both faba bean
(Fig. 2) and chickpea (Lake and Sadras, 2014) have broad critical
periods and yield responds to shading early in the season in contrast to
cereals and canola (Fischer, 1985; Kiniry and Ritchie, 1985; Savin and
Slafer, 1991; Arisnabarreta and Miralles, 2008; Estrada-Campuzano
et al., 2008; Oosterom and Hammer, 2008; Cerrudo et al., 2013;
Mahadevan et al., 2016; Kirkegaard et al., 2018; Ali et al., 2019).
The reasons for a broader sensitivity period in pulses are unknown.
In pulses, the time lag between flowering and pod set can be extended
under stressful conditions (Mwanamwenge et al., 1998, 1999; Berger
et al., 2004). Reduction in yield with early-season stress maybe par-
tially related to shade induced reduction in nodulation, and reduced
nitrogenase activity (Rao and Mittra, 1988; Matamoros et al., 1999).
The critical period was not affected by the different evapo-
transpiration environments emphasising the intrinsic crop-specific re-
sponsiveness to stress Yield response to shading was most severe from
flowering to pod emergence, conforming with other indeterminate
species including soybean, cool-season legumes and canola (Board and
Tan, 1995; Jiang and Egli, 1995; Sandaña and Calderini, 2012; Lake
and Sadras, 2014; Kirkegaard et al., 2018). This period coincides with
both vegetative and reproductive growth, maximum nitrogen fixation
rate and the largest number of flowers and embryos that are yet to
reach the final stage in seed abortion (Munier-Jolain et al., 1993; Ney
et al., 1993). In common with other species and consistent with the
theory of yield determination in grain crops (Sadras, 2007; Sadras and
Slafer, 2012; Slafer et al., 2014), seed number mediated yield response
to shading. Similarly to chickpea, lupin and field pea, pod number was
the primary determinant of seed number when shaded before flow-
ering, with some influence of seeds per pod when shaded after flow-
ering (Sandaña and Calderini, 2012; Lake and Sadras, 2014). The in-
determinate grain legumes thus appear to differ with canola, for which
seeds per pod were more responsive to shading than pod number

4
L. Lake, et al.
Fig. 2. Effect of timing of shading on (a) yield, (b) seed number and (c) seed
size of faba bean from Tamworth 2017 (triangle), Breeza 2018 (hexagon),
Roseworthy 2017 (square), Roseworthy 2018 (circle) and Valdivia 2018 (dia-
mond). Values are normalised relative to controls. Varieties are pooled. Open
symbols show no effect of shading relative to control, and closed symbols in-
dicate difference at P < 0.05. The lines are a combination of polynomial re-
gression where shading treatment differs from controls, and the horizontal
segments are averages where they are not. Error bars are ± S.E and are not
shown when smaller than symbol. The phenological scale is based on the un-
shaded controls.
(Kirkegaard et al., 2018).
Crops can partially compensate reductions in seed number with
increased seed size with some crops fully compensating such as canola
(Labra et al., 2017). In our study, faba bean slightly increased seed size
when shaded before flowering, and again after pod set (Fig. 2c). In
comparison, chickpea increased seed size in response to shading after,
but not before flowering (Lake and Sadras, 2014). Faba bean was also
less sensitive to reductions in seed per pod later in the season compared
with chickpea (Lake and Sadras, 2014). This could indicate that faba
bean reaches the final stage in seed abortion earlier than chickpea, or it
may be a result of faba bean eventually reaching its terminal flores-
cence, or a combination (Stoddard, 1993; Munier-Jolain et al., 1998;
Walley et al., 2007). Specific causes for yield sensitivity around the
critical period deserve further research.
Characterisation of the critical period can aid in crop breeding by i)
increasing the likelihood of successful selection strategies, ii) increasing
the efficiency of screening by narrowing the focus to critical physiolo-
gically relevant stages, and will also aid in improved agronomy and
stress mitigation practices (Andrade et al., 2005; Sandaña and
Calderini, 2012; Cerrudo et al., 2013).
5
Field Crops Research 241 (2019) 107575
Fig. 3. Effect of timing of shading on (a) pod number and (b) seed per pod of
faba bean from Tamworth 2017 (triangle), Breeza 2018 (hexagon), Roseworthy
2017 (square), and Roseworthy 2018 (circle) and Chile 2018 (diamond). Values
are normalised relative to controls. Varieties are pooled. Open symbols show no
effect of shading relative to control, and closed symbols indicate difference at
P < 0.05. The lines are a combination of polynomial regression where shading
treatment differs from controls, and the horizontal segments are averages where
they are not. The phenological scale is based on the unshaded controls. Error
bars are ± S.E and are not shown when smaller than symbol.
Acknowledgements
We thank the technical assistance of Penny Roberts, John Nairn,
Phil Rundle, Tanja Lenz, Jerman Carrasco, Han Chow, Annabel O’Dea,
Justyn Thompson, Jose Fernandez, Matt Grinter, Michael Nowland and
Peter Sanson, and the financial support by the Grains Research and
Development Corporation-SARDI bilateral (project DAS00166_BA) and
the Chilean Technical and Scientific Research Council (CONICYT)
Project FONDECYT 1170913 competitive grant.
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