Immunity and other defenses in pea aphids, Acyrthosiphon pisum
Effect of harvest time on yield and pre-harvest quality of semi-leafless grain peas ( Pisum sativum...
Transcript of Effect of harvest time on yield and pre-harvest quality of semi-leafless grain peas ( Pisum sativum...
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Field Crops Research 100 (2007) 1–9
Effect of harvest time on yield and pre-harvest quality of semi-leafless
grain peas (Pisum sativum L.) as whole-crop forage
Giorgio Borreani a,*, Pier Giorgio Peiretti b, Ernesto Tabacco a
a Dipartimento di Agronomia, Selvicoltura e Gestione del Territorio, Universita di Torino, via L. da Vinci 44, 10095 Grugliasco (TO), Italyb Istituto di Scienze delle Produzioni Alimentari, CNR, via L. da Vinci 44, 10095 Grugliasco (TO), Italy
Received 3 November 2005; received in revised form 19 April 2006; accepted 19 April 2006
Abstract
The aim of the work was to study changes in the yield and nutritional characteristics of whole crop semi-leafless field pea over two growing
seasons in the Po plain, Italy. Samples of two cultivars (Baccara and Sidney) were collected from flowering to grain maturity. The
developmental stage, yield, dry matter (DM) content, crude protein (CP), neutral detergent fibre (NDF), acid detergent fibre (ADF), acid
detergent lignin (ADL), starch, water soluble carbohydrates (WSC), gross energy (GE), organic matter digestibility (OMD) and the net energy
for lactation (NEL) were determined at each harvest. The forage characteristics were regressed on the growing degree days (GDD) with 4.4 8Cas the base temperature. The DM yield increased with advancing maturity from 0.5 to 8.91 Mg ha�1, while the CP decreased from 261 to
159 g kg�1 DM. During the whole growth cycle the GE, OMD, NEL and milk forage units (milk FU) were almost steady. No differences were
observed between the cultivars for any of the measured parameters. At grain maturity, the crop produced over 4.0 Mg ha�1 DM of grain. The
CP, starch and WSC of the grain did not show any differences between the cultivars or years. The data showed that the nutritive quality of the
forage of the semi-leafless grain pea harvested as a whole crop for ensiling purposes did not diminish with maturity and could help improve the
self-sufficiency of dairy farms, in terms of home-grown protein forages.
# 2006 Elsevier B.V. All rights reserved.
Keywords: Growing degree days; Growth stage; DM yield; OM digestibility; NDF; Peas
1. Introduction
The European bans on feeding fish or animal protein meals
to livestock have highlighted the huge deficit in plant protein
in the EU (>75% in 2000), and have led to considerable
pressure on livestock farmers to produce home-grown
alternative forages (Wilkins and Jones, 2000), that are high
in protein and energy. Peas (Pisum sativum L.) are mainly
used for grain production throughout the world, however, this
species is sometime used alone or in combination with spring
cereals as whole-crop forage (Faulkner, 1985; Mustafa et al.,
2002). Pea seed contain a moderate amount of high quality
protein and a high level of starch, thus making this crop a
unique dual-purpose feed, rich in both energy and protein
(Cousin, 1997). Peas can offer farmers a great deal of
* Corresponding author. Tel.: +39 011 6708783; fax: +39 011 6708798.
E-mail address: [email protected] (G. Borreani).
0378-4290/$ – see front matter # 2006 Elsevier B.V. All rights reserved.
doi:10.1016/j.fcr.2006.04.007
flexibility in their cropping rotations due to their quick growth
in a short period providing forage of a high nutritive value and
palatability (Koivisto et al., 2003). Due to their relatively high
moisture content, peas are rarely harvested for hay and they
are most often ensiled as a whole crop to provide both protein
and starch sources for livestock (Mustafa and Seguin, 2004),
which allows for a minimal feeding of purchased concentrates
without adversely affecting milk yield or quality (Adesogan
et al., 2004). Mustafa et al. (2000) reported that whole pea
silages, harvested at full pod stage, can successfully replace
barley or alfalfa silage as a forage source for high yielding
dairy cows in early lactation, while Børsting and Weisbjerg
(2002) found that feeding pea silage harvested with
undeveloped seeds to dairy cows led to a lower dry matter
(DM) intake, milk yield, and milk protein yield in comparison
to perennial ryegrass silage made with a comparable crude
protein content. Date of harvest and hence stage of maturity at
which crops are harvested will greatly affect forage yield and
G. Borreani et al. / Field Crops Research 100 (2007) 1–92
quality. Several studies have reported that delaying harvesting
of pea past the flowering stage may result in significantly
greater forage DM yields (Aman and Graham, 1987; Mustafa
and Seguin, 2004). Reduction in forage and seed yield have
often been attributed to lodging, which is particularly evident
in the leafed varieties, while the semi-leafless phenotypes are
less susceptible (Koivisto et al., 2003; Uzun et al., 2005). With
grasses and leguminous forages quality forage can be
obtained by management strategies that are directed towards
cutting when the leaf to stem ratio is high (Marten et al., 1988;
Nelson and Moser, 1994). When pulses are to be used as
whole-crop forage the management strategies differ because
the grain and pod yields are also important (Salawu et al.,
2001). Selection of an optimal harvest date will require a
better understanding of the chemical changes that occur
during maturation and of the feeding value of the crop at
different stages of development (Aman and Graham, 1987).
Nutritive value has been related to accumulated growing
degree days (GDD), crop age in days, leaf to stem ratio, stand
height and codified morphological stage (Nelson and Moser,
1994). The GDD, involving the sum of daily mean
temperature above a given base temperature, is widely used
with peas to predict relative maturity during the growing
season (Bourgeois et al., 2000; Mikkelsen, 1981). A base
temperature from 0 to 5.0 8C was generally adequate for all
cultivars to compute growing degree days between sowing
and maturity, although a base temperature of 4.4 8C was often
reported to be the most satisfactory for green pea (Bourgeois
et al., 2000).
Little information can be found in the literature on the
changes in yield potential and nutritional quality during the
growth cycle of peas sown in spring and grown to be ensiled
as a whole-crop in southern Europe. The objective of this
study was to provide information on the effect of harvest
time on the pre-harvest nutritional quality and DM yield of
two cultivar of semi-leafless peas over a 2-year period in the
Po plain in Northern Italy.
2. Materials and methods
2.1. Plant material and environmental conditions
The research was carried out in the 2001 and 2002 growing
seasons in the Western Po plain near Torino (448500N, 78400E,
altitude 232 m above sea level, annual mean temperature
11.7 8C, and annual average rainfall 739 mm) on a sandy-
loam textured alluvium soil with a pH measured in water of
7.6. The sand, silt, and clay contents of the soil were 470, 440,
and 90 g kg�1, respectively at 0–30 cm depth. The organic C
was 11.8 g kg�1 and organic N was 1.45 g kg�1. Two semi-
leafless pea cultivars (Baccara and Sydney) were selected as
they were among the best known commercial lines adapted to
the area, at the time of the initiation of the study (Battini et al.,
2001). Stands were sown at 230 kg ha�1 of naked viable seed
on 21 March in 2001 and at 340 kg ha�1 on 28 February in
2002 to enable better competition with weeds. No irrigation or
fertiliser was applied after sowing. Weeds were controlled by
a post-emergence treatment using 0.9 kg a.i. ha�1 of benta-
zone in 2001. Temperature and rainfall data were collected
from a weather station approximately 60 m from the
experiment site. Herbage samples for the yield and quality
measurements were collected with a Haldrup forage plot
harvester on 12 m2 randomly located subplots with two
replicates in 30 m � 45 m main plots (two plots per cultivar).
Forage was cut to a 3–4 cm stubble height. Herbage samples
were collected four to five times at progressive morphological
stages from flowering to late seed pod stage from the middle of
May to the end of June.
At each sampling time, when pods were present, sub-
samples of 30 plants were divided into seeds and other plant
components (tendrils, stipules, stems, inflorescences and
pods), to determine the grain contribution to the DM yield.
Crops were also harvested at grain maturity (beginning of
July) to evaluate the grain and straw production and quality.
A complete nutritive analysis of the straw was only made in
2002 in order to better characterise the role of this crop
component for animal nutrition purposes.
2.2. Chemical analysis
Chemical analyses were performed on the whole-plant,
the grain and the straw samples. The whole-plant and straw
samples were chopped to a length of 20–30 mm. All samples
were immediately dried in a forced-draft oven at 60 8C, air
equilibrated, weighed, ground in a Cyclotec mill (Tecator,
Herndon, VA, USA) to pass a 1 mm screen, and stored for
qualitative analyses.
The dried samples were analysed to determine the total N,
according to the Dumas method, using a Nitrogen analyser
Micro-N (Elementar, Hanau, Germany). The crude protein
(CP) (total N x 6.25), ash by ignition to 550 8C, neutral
detergent fibre (NDF), acid detergent fibre (ADF), and acid
detergent lignin (ADL) were determined according to
Robertson and Van Soest (1981), while the sugar and starch
were determined as described in the subsequent paragraph.
Gross energy (GE) was determined using an adiabatic
calorimeter bomb (IKA C7000, Staufen, Germany), and the
organic matter digestibility (OMD) according to the two-
stage rumen fluid technique (Tilley and Terry, 1963). The
OMD values were expressed in vivo using the Goldman et al.
(1987) regression equation. Net energy, expressed as both
the net energy for lactation (NEL) and milk forage unit (milk
FU), was calculated by inserting the observed values of the
OMD and GE into the equations proposed by Andrieu and
Demarquilly (1987).
2.3. Sugar extraction and starch hydrolysis
measurements
Sugar and starch were determined using the methods
described by Rong et al. (1996) with some modifications.
G. Borreani et al. / Field Crops Research 100 (2007) 1–9 3
Table 1
Monthly and annual mean temperatures and accumulated precipitation for
the study period and the long term average
Month Temperature (8C) Rain (mm)
2001 2002 Mean
1976–2002
2001 2002 Mean
1976–2002
January 1.5 �1.7 0.7 16 6 36
February 4.8 4.1 3.3 16 149 33
March 8.7 9.4 7.7 68 55 52
April 11.1 12.0 10.9 10 64 96
May 17.9 16.2 16.0 183 153 112
June 20.1 22.1 19.6 8 81 74
July 22.3 21.6 22.0 25 129 39
Annual 12.3 12.3 11.7 480 1191 739
Approximately 8 mg of grains or 20 mg of whole plant were
rinsed four times with 80% (v/v) ethanol, using 1.0 mL
ethanol per rinse, centrifuged, and the supernatant decanted
into a graduated test tube. An anthrone reagent was added to
an aliquot containing up to 200 mg of free sugar, boiled for
20 min, and then cooled to room temperature in a cold water
bath. Absorbance was determined at 625 nm with a Unicam
Helios g spectrophotometer (Unicam, Cambridge, UK).
Total sugar contents were expressed on the basis of glucose
equivalent standards containing 0, 50, 75, 100, 150, and
200 mg glucose mL�1 80% ethanol.
The tube containing the ethanol-insoluble pellet was
placed in a 55 8C oven for 24 h to evaporate any residual
ethanol before starch hydrolysis. Distilled deionized water
(0.5 mL) was added to each tube and the starch was
gelatinized by boiling the samples for 10 min. After cooling
to room temperature, 2 U amyloglucosidase (Sigma Chemical
product A3514) and 80 U a-amylase (Sigma Chemical
product A2643) were added to each tube and incubated at
55 8C for 24 h in a 200 M acetate buffer (pH 5.0). At the end of
the incubation period, the samples were centrifuged at
16,000 � g for 10 min. The glucose released from the starch
hydrolysis was determined with a glucose Trinder reagent
(Sigma Chemical Diagnostic kit No. 315). Absorbance was
read at 505 nm and the results were compared with glucose
standards containing 0, 0.5, 1, 2.5, and 5 mg glucose mL�1
distilled deionized water. The starch concentration was
calculated by multiplying the glucose concentration by 0.9.
2.4. Statistical analysis
The data were analysed over the years and harvest maturity
through regression analysis and via an analysis of variance
(ANOVA), using the Statistical Package for Social Science (v
11.5, SPSS Inc., Chicago, IL, USA). All the chemical and
nutritional parameter analyses were performed in duplicate
and averaged for the statistical analyses. The resulting data
were averaged over the sub-plot replicates before regression
analysis and used as individual observations for ANOVA. The
data were regressed on the growing degree days (GDD) with a
4.4 8C base temperature (Mikkelsen, 1981) and on the age in
days from the seeding date, as independent variables. The
accumulated GDD was calculated using the following
formula for each year:P
{[(Tmax + Tmin)/2] � 4.4 8C};
where Tmax and Tmin are the daily maximum and minimum
temperatures, respectively, in degree Celsius. The average
temperature [(Tmax + Tmin)/2] was set equal to 4.4 8C if less
than 4.4 8C (McMaster and Wilhelm, 1997).
Linear and quadratic regressions were compared using
the Draper and Smith (1998) stepwise selection procedure to
select the best regression model at the 0.05 probability level.
The best equation for each parameter was selected using
coefficient of determination and root mean square error
(RMSE). All the determination coefficients (R2) reported in
this paper were adjusted for degrees of freedom. The
MANOVA analysis of covariance was used to verify the
equivalence of the equations for the cultivars and years. This
procedure allowed one to verify the hypothesis of
parallelism of the regression equations and, when this
was not rejected, the analysis of covariance was used to
evaluate the differences between the adjusted means. When
no differences were found a single regression line was
determined for the pooled data.
The chemical compositional data were analysed via
ANOVA, with their significance reported at a 0.05
probability level using the general linear model of SPSS.
3. Results
3.1. Weather data
The monthly means of air temperature and accumulated
rain, as well as the annual mean temperature and the annual
accumulated rain of the two experimental years from
January to July and the 27-year mean values are reported in
Table 1. The 2 years were characterised by mean
temperatures that were higher than the 27-year values,
especially in February and March. The accumulated GDD
were lower in 2002 than in 2001, due to the earlier seeding
date (28 February versus 21 March).
3.2. Morphological stage and DM yield
The effects of the cultivar and harvesting date on the DM
yield and grain proportion are shown in Table 2. The stages
considered before grain maturity were: late bud, 17 May
(Cut 1); end of flowering, 31 May (Cut 2); beginning of pod
filling, 7 June (Cut 3); advanced pod filling, 14 June (Cut 4),
and beginning of ripening, 21 June (Cut 5) in 2001; and 20%
of flowering, 15 May (Cut 1); beginning of pod filling, 3 June
(Cut 2); advanced pod filling, 13 June (Cut 3); and mid-
ripening, 20 June (Cut 4) in 2002. The physiological
maturity of the seeds was reached in 106 and 123 days from
seeding in 2001 and 2002, respectively, corresponding to
1259 and 1309 8C of GDD.
G. Borreani et al. / Field Crops Research 100 (2007) 1–94
Tab
le2
Gro
win
gd
egre
ed
ays
(GD
D),
DM
yie
ldan
dg
rain
pro
port
ion
of
the
two
cult
ivar
sin
20
01
and
20
02
Yea
r
20
01
20
02
Cu
ltiv
arC
ut
1C
ut
2C
ut
3C
ut
4C
ut
5G
rain
mat
uri
tyC
ut
1C
ut
2C
ut
3C
ut
4G
rain
mat
uri
ty
Dat
e1
7M
ay3
1M
ay7
Jun
e1
4Ju
ne
21
Jun
e5
July
15
May
3Ju
ne
13
Jun
e2
0Ju
ne
1Ju
ly
Day
sfr
om
seed
ing
57
71
78
85
92
10
67
69
51
05
11
21
23
Sta
ge
of
mat
uri
tyL
ate
bu
dE
nd
of
flow
erin
g
Beg
inn
ing
po
dfi
ll
En
dp
od
fill
Beg
inn
ing
rip
enin
g
Gra
in
mat
uri
ty
20
%
flow
erin
g
Beg
inn
ing
po
dfi
ll
En
dp
od
fill
Mid
rip
enin
g
Gra
in
mat
uri
ty
GD
D(4
.48C
)4
85
70
78
02
90
01
00
11
25
95
39
80
99
53
11
01
13
09
DM
yie
ld(M
gh
a�1)
Bac
cara
0.5
03
.66
5.4
67
.21
7.2
78
.69
1.2
84
.26
5.2
06
.18
6.6
2
Sid
ney
0.6
63
.64
4.6
57
.05
7.8
58
.91
1.5
14
.66
6.1
06
.36
7.4
1
Gra
inp
rop
ort
ion
(%D
M)
Bac
cara
00
11
25
48
48
03
94
56
46
1
Sid
ney
00
72
85
04
80
31
50
65
64
The different seeding rates in the 2 years resulted in plant
densities of 51 and 49 plants m�2 and 98 and 96 plants m�2,
for Baccara and Sidney in 2001 and 2002, respectively. The
DM yield increased with stage of maturity from 0.50 to
8.69 Mg ha�1 and from 0.66 to 8.91 Mg ha�1 in 2001 for
Baccara and Sidney, respectively and from 1.28 to
6.62 Mg ha�1 and from 1.51 to 7.41 Mg ha�1 in 2002 for
Baccara and Sidney, respectively. These increases were
obtained in 50 and 48 growing days, corresponding to 774
and 770 8C of GDD for 2001 and 2002, respectively. The
grain contribution to the DM yield was similar for the two
cultivars and reached values of 48% of the total DM in 2001
for both cultivars, and values of 61 and 64% in 2002, for
Baccara and Sidney, respectively.
3.3. Variation in DM content and nutritional quality of
the whole-plant during the growth cycle
The GDD was always a better descriptor of the change in
forage quality over the growth cycle from the beginning of
flowering to mid-ripening than the age in days from seeding.
For this reason, the following results are shown in relation to
the GDD only (Figs. 1–4).
The DM content of the herbage was low for early harvests
before flowering, with values of around 110 g kg�1, but
reachedvaluesof 191and221at advanced podfilling, and then
rapidly increased duringripening of the seed pod(Fig. 1a). The
ash content decreased with maturity from 115 to 72 g kg�1
DM for 2001 and from 84 to 67 g kg�1 DM for 2002, without
any differences between the cultivars. The CP content of the
whole plant decreased with advancing maturity with different
trends for the 2 years, both with high coefficients of
determination. No differences were observed between the
cultivars. After the beginning of pod filling, the CP was almost
constant at around a mean value of 170 g kg�1 DM for both
years (Fig. 1b). The WSC showed different trends for the years
in relation to the GDD. The values increased from 41 to
88 g kg�1 DM till the fourth cut and then decreased to
47 g kg�1 DM in the fifth cut in 2001, while it decreased with
increasing GDD from 115 to 15 g kg�1 DM, in 2002 (Fig. 2a).
The starch was directly related to the GDD with a single
quadratic regression line over the years and cultivars with
coefficient of determination greater than 0.90. The starch
values ranged from 19 to 249 g kg�1 DM with plant
development (Fig. 2b). The NDF content increased with the
same trend over the years and for the cultivars (Fig. 2c) with a
coefficient of determination of 0.904. TheADFand ADLwere
almost constant over the years for the cultivars with a mean
value of 235� 18.5 g kg�1 DM and 35.1 � 5.4 g kg�1 DM,
respectively. The GE and OMD of the herbage did not show
any significant relationship with the GDD and were almost
constant over the years and for the cultivars with a mean value
of 17.5 � 0.35 MJ kg�1 DM and of 806� 17.4 g kg�1 OM,
respectively. Consequently both the NEL and milk FU were
almost constant with a mean value of 6.80 � 0.24 MJ kg�1
and 0.955� 0.034 milk FU kg�1, respectively.
G. Borreani et al. / Field Crops Research 100 (2007) 1–9 5
Fig. 1. Dry matter (DM) (a) and crude protein (CP) (b) content of whole plant peas as a function of the growing degree days (GDD); filled symbols, 2001; empty
symbols, 2002. When no differences were found a single regression line was determined for the pooled data: (a) DM = 0.000847 GDD2 � 1.024 GDD + 445;
R2 = 0.920; RMSE = 19.7; (b) CP(2001) = 0.000172 GDD2 � 0.426 GDD + 427; R2 = 0.907; RMSE = 8.5; CP(2002) = 0.0878 GDD + 263; R2 = 0.761;
RMSE = 10.4.
Fig. 2. Water soluble carbohydrate (WSC) (a), starch (b) and NDF (c) contents of whole plant peas as a function of the growing degree days (GDD); filled
symbols, 2001; empty symbols, 2002. When no differences were found a single regression line was determined for the pooled data: (a) WSC(2001) = �0.000433
GDD2 + 0.693 GDD � 200; R2 = 0.595; RMSE = 10.7; WSC(2002) = �0.000486 GDD2 + 0.646 GDD � 100; R2 = 0.950; RMSE = 8.1; (b) Starch(2001–
2002) = 0.000186 GDD2 + 16.36; R2 = 0.914; RMSE = 16.9; (c) NDF(2001–2002) = 0.000504 GDD2 � 0.461 GDD + 409; R2 = 0.904; RMSE = 21.3.
3.4. Evolution of nutritional quality of plant
components during crop growth
The changes in the straw CP and NDF contents during
the growth cycle are reported in Fig. 3. The CP content
decreased with advancing maturity, following different
trends over the 2 years. In the period from the end of
flowering to the beginning of the pod filling stage (from
Fig. 3. Crude protein (CP) (a) and neutral detergent fibre (NDF) (b) of pea straw a
symbols, 2002. When no differences were found a single regression line was dete
RMSE = 12.7; CP(2002) = 0.000332 GDD2 � 0.904 GDD + 703; R2 = 0.924; R
R2 = 0.923; RMSE = 21.8.
700 to 800 8C of accumulated GDD) the CP content was
slightly lower than 200 g kg�1 DM, and decreased to a
mean value of 141 g kg�1 DM in 2001 and 91 g kg�1 DM
in 2002 at grain harvest (GDD from 1250 to 1300 8C)
(Fig. 3a). The NDF increased from values around
400 g kg�1 DM to higher than 500 g kg�1 DM in 2001
and 600 g kg�1 DM in 2002, respectively, at grain
harvesting (Fig. 3b).
s a function of the growing degree days (GDD); filled symbols, 2001; empty
rmined for the pooled data: (a) CP(2001) = 0.0861 GDD + 247; R2 = 0.684;
MSE = 11.1; (b) NDF(2001–2002) = 0.000494 GDD2 – 0.656 GDD + 602;
G. Borreani et al. / Field Crops Research 100 (2007) 1–96
Fig. 4. Crude protein (CP) (a), water soluble carbohydrate (WSC) (b) and starch (c) contents of pea grain as a function of the growing degree days (GDD); filled
symbols, 2001; empty symbols, 2002. When no differences were found a single regression line was determined for the pooled data: (a) CP(2001–2002) = 0.000634
GDD2 � 1.414 GDD + 1010; R2 = 0.700; RMSE = 12.4; (b) WSC(2001–2002) = 0.000911 GDD2 � 2.136 GDD + 1262; R2 = 0.973; RMSE = 6.9; (c)
Starch(2001–2002) = �0.00322 GDD2 + 7.374 GDD � 3597; R2 = 0.876; RMSE = 44.7.
The changes in grain CP, WSC and starch content during
the growth cycle are reported in Fig. 4. The crude protein
content of the grain during the filling of the seeds slightly
decreased from 286 g kg�1 DM in the first phase to a mean
value of 234 g kg�1 DM in the following maturation phases
with no differences between the years or cultivars (Fig. 4a).
The WSC content decreased faster with advancing maturity
to values of around 20 g kg�1 DM (Fig. 4b). The decrease in
WSC was coupled to an increase in starch accumulation in
the seed tissue that increased from around 250 g kg�1 DM to
values of 556 g kg�1 DM (Fig. 4c).
3.5. Nutritional quality of grain and straw at grain
maturity
The DM yield and composition of the grain and of the
straw at the end of the cycle over the years and for the
cultivars are reported in Table 3. The stands produced over
Table 3
Dry matter (DM) yield, crude protein (CP), starch content and water soluble carboh
for straw for the two cultivars at grain maturity, in 2001 and 2002
Year
2001 2002
Baccaraa Sidney Baccara
Grain
DM yield (Mg ha�1) 4.21 4.28 4.06
CP (g kg�1 DM) 249 218 246
Starch (g kg�1 DM) 553 530 577
WSC (g kg�1 DM) 20 20 22
Straw
DM yield (Mg ha�1) 4.48 4.63 2.57
CP (g kg�1 DM) 131 151 96
NDF (g kg�1 DM) 532 542 586
a Cultivar.b C: cultivar effect; Y: year effect; S.E.M.: standard error of the mean.c NS: not significant, P > 0.05.* P < 0.05
** P < 0.01.*** P < 0.001.
4.0 Mg DM ha�1 of grain for the two cultivars and years.
The crude protein, starch and WSC did not show any
differences between the cultivars or years.
Straw DM yield in 2001 was about double that of 2002,
with no differences between the cultivars. The crude protein
of the straw was higher in 2001 than in 2002 (141 g kg�1 DM
versus 91 g kg�1 DM) with slight differences between
cultivars in 2001, while the NDF was higher in 2002 and
for the Sidney cultivar. A complete nutritive analysis of the
straw was performed in 2002. No significant differences were
found between the cultivars except for the ADL values
(94 g kg�1 versus 106 g kg�1 DM for Baccara and Sidney,
respectively). The OMD, GE, NEL, and milk FU mean values
were 471� 33 g kg�1 OM, 16.5 � 0.4 MJ kg�1 DM,
3.33 � 0.26 MJ kg�1 DM, 0.47 � 0.04 kg�1 DM, respec-
tively. The residual starch and WSC values were
14.9 � 9.5 g kg�1 DM and 5.9� 1.5 g kg�1 DM, respec-
tively.
ydrate (WSC) for grain and DM yield, CP and neutral detergent fibre (NDF)
Yb C Y � C S.E.M.
Sidney
4.74 NSc NS NS 0.67
241 NS NS NS 15.7
566 NS NS NS 56.3
20 NS NS NS 2.53
2.67 ** NS NS 0.48
86 *** NS * 9.95
627 ** * NS 19.9
G. Borreani et al. / Field Crops Research 100 (2007) 1–9 7
4. Discussion
By sowing the crop in March rather than the fall,
excellent stand establishment was obtained, and the risk of
winter chill damage was minimized. The GDD was found to
give a more reliable description of the evolution of the
agronomic and nutritional parameters and to be more
independent of the sowing time than the age in days from
seeding date. This has also been observed in lucerne by Fick
et al. (1994) and in sainfoin by Borreani et al. (2003).
The crop produced a high DM yield in the 13 or 16-week
growing cycles from seeding to harvesting for the two years.
This short-term catch-crop harvested as silage would allow a
subsequent maize or soybean crop to be grown in the same
year in the Po plain environment (Tabacco, personal
communication). A DM yield higher than 5.0 Mg ha�1
was reached at the beginning of pod fill; Trevino et al. (1987)
observed halved yields at the same stage of growth in Spain,
while similar yields were observed by Mustafa and Seguin
(2004) between 8 and 10 weeks after seeding in the colder
Canadian environment, both for peas cultivated on their own
or as a pea-cereal mixture. In the UK environment, Fraser
et al. (2001) obtained 5.4 Mg ha�1 at 10 weeks from seeding
and 6.2 Mg ha�1 at 12 weeks using cv. Magnus. In our
experiment the DM yield of the crop continued to increase in
both years throughout the whole growth cycle. Sheldrick
et al. (1995) observed that delaying harvesting beyond the
flat pod stage resulted in a limited increase in DM yield
because of the senescence of the lower leaves and stems.
This difference can be partially explained by the better
standing ability of the semi-leafless pea lines than the leafed
lines, which leads to a delay of or prevents lodging of the
crop. Together with the higher DM yield, a more mature
plant can also have a higher DM content, possibly
eliminating the need to wilt the crop prior to harvest and,
thereby, potentially reducing the field losses (Fraser et al.,
2001). However, it is likely that allowing the crop to mature
to a late stage before harvesting may cause a detrimental
effect on the crude protein, fibre content and digestibility of
the crop (Van Soest, 1994). It is known that the
morphological development of the plant is the factor that
most affects the nutritive value of several grass and legume
forages (Van Soest, 1994). The large decline in the protein
and nutritive value of about 40% is attributed to the
interactive effects of the increase in fibre and lignin in the
tissues, which is more marked in stem tissue than in leaf
tissue, and to the different ratios between the plant tissue
components (Van Soest, 1994). This interactive effect is less
evident in crops where the grain contribution to the DM
yield is high (Salawu et al., 2001). In cereals (Kennelly and
Weinberg, 2003) and pulses (Panciera et al., 2003), utilized
as whole-crops, grain contribution to the total harvested DM
can reach 50% or more. In this experiment, the grain
contribution increased with advancing maturity and reached
50 and 65% after 92 and 112 days from sowing for 2001 and
2002, respectively. Koivisto et al. (2003), in the UK
environment, observed that pods of grain-type cultivars
accounted for 25% of the total mass of the plant at 80 days
after sowing. At a comparable harvesting date we observed
similar values with pods accounting for 26 and 35% of the
total plant mass, for the two experimental years.
The crop yielded from 0.9 to 1.3 Mg ha�1 of protein from
the beginning to the end of pod filling (from 10 to 14 weeks
after sowing) when harvested for silage purposes. These
values are slightly higher than those observed by Mustafa
and Seguin (2004) who harvested around 0.8 Mg ha�1 of
protein in a whole-crop pea at 10 weeks from seeding (pod
filling stage).
The NDF of the whole crop increased over the growth
cycle because, as the plant grows, the need for structural
tissue increases, and the NDF of the straw increased from
400 to more than 600 g kg�1 DM. The ADF, ADL and the
nutritive value of the whole crop pea expressed as GE, OMD
and NEL were constant over the whole growth cycle. This
can be explained by a decrease in the digestibility of the pea
straw with advancing maturity which is balanced by an
increase in grain proportion. These results are in agreement
with Mustafa and Seguin (2004) and Salawu et al. (2001),
who found that the in vitro OMD in the DM of pea/wheat bi-
crops did not appear to be significantly influenced by the
stage of maturity at harvesting. Fraser et al. (2005) also
observed no drop in digestibility for lupin (Lupinus albus)
silages harvested at four different stages of growth. Similar
results have been reported for whole maize silage where in
vitro DM digestibility was almost constant from late
flowering till advanced dough stage, due to the increase
in grain proportion (Johnson et al., 1999), while, in contrast,
Crovetto et al. (1998) observed a decrease in the in vivo
digestibility of whole-crop wheat silage and in the net
energy for lactation with advancing stage of maturity from
boot stage onwards.
In this experiment pea seeds contained an average CP of
243 g kg�1 DM that remained almost constant for the whole
growth cycle. Daveby et al. (1993) observed the same
pattern for three Swedish pea cultivars. Analyses of 1100
samples compiled over 4 years from throughout western
Canada confirmed a feed pea CP level of 239 g kg�1 DM
(Fleury, 2004). The WSC content was higher in the earlier
stages of seed development and rapidly decreased after
900 8C of GDD, corresponding to the advanced pod fill, as a
consequence of starch synthesis in the seed. Haeder (1989)
reported that synthesis of amylose and amylopectin, the two
main components of starch, increased rapidly after the first
third of seed DM is produced. The starch values of pea seeds
at harvest averaged 556 g kg�1 DM. This is higher than the
average starch values of 453 g kg�1 DM (Cerioli et al.,
1998) and 501 g kg�1 DM (Grosjean et al., 1999) measured
in extensive European surveys.
Pea straw could be a good feed alternative in beef diets,
due to its higher feed value than wheat or barley straw
(Cherriere, 2001). Pea straw has been reported to contain, on
average, 64 g kg�1 DM of CP, 695 g kg�1 DM of NDF,
G. Borreani et al. / Field Crops Research 100 (2007) 1–98
while at grain maturity we found higher value for CP and
lower values of NDF. Alternatively the straws could be
incorporated into the soil, and could contribute from 23 to
70 kg N ha�1 for the subsequent crop.
5. Conclusions
Semi-leafless grain peas seeded at the end of winter and
harvested from the middle of May to the end of June can give
high yields, providing good quality forage for ruminants and
may improve the self-sufficiency of dairy farms, in terms of
home-grown protein forages. The high DM yields,
associated with a low NDF content and a high OM
digestibility, suggest that whole crop peas should be
harvested and ensiled from the advanced pod filling stage
onwards, since semi-leafless grain peas did not show a
decline in the quality of forage with maturity.
Acknowledgements
The authors wish to thank Mario Gilardi and Mauro Gilli
(Dipartimento di Agronomia, Selvicoltura e Gestione del
Territorio) for the technical assistance in the field. Financial
support for this work was provided by the Consiglio
Nazionale delle Ricerche and by the MURST Funding
Program ‘‘60%’’. All the authors contributed equally to the
work described in this paper.
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