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ABSTRACT ry pea has long been known as an excellent source
of protein in human and livestock diet (Hood-Niefer
Pea (Pisum sativum L.) has long been an important component
et al., 2012), and plays an important role in sustain-
of the human diet, providing an excellent source of protein. In
able cropping systems worldwide. In the U.S. northern Great
addition to its protein, pea starch, especially resistant starch
Plains (NGP), a shift toward a cerealdry pea cropping system
(RS), has received an extensive attention in food industries in
has been a major trend of the agricultural industry (Long et
recent years. We evaluated nine pea cultivars varying in cotyle-
al., 2014). Intensification and diversification of the cropping
don color, grain weight, maturity group, and phenology planted
systems with pulse crops, mainly dry pea, in this region has
at five locations with diverse climatic conditions across Montana
brought benefits to growers by creating an immediate economic
in 2013 and 2014 to assess genetic and environmental factors
return, providing typical rotational benefit, and offering N
affecting their yield, protein, RS, and total starch (TS). Grain
credit for the subsequent crop yield and grain quality (Chen et
yield varied from 982 to 5951 kg ha1, RS content ranged from 5
al., 2012; Ito et al., 2016).
to 53 g kg1, and protein from 159 to 251 g kg1. Statistical anal-
Some efforts have been made to quantify yield variation
ysis showed that environment was the most important driving
of dry pea across NGP. For example, Ito et al. (2016) evalu-
factor in grain yield, protein, and TS determination whereas RS
ated adaptation and yield stability of dry pea cultivars across
content was mainly determined by cultivar. Drought at all phe-
Montana from 2009 to 2011. They reported that a large por-
nological stages reduced pea yield and different cultivars tended
tion of the total variation of yield was explained by the envi-
to respond differently. Yield was positively correlated with pro-
ronment (E). In that study, Delta, Majoret, and Cruiser were
tein, implying a potential to select/breed a cultivar with higher
found as suitable cultivars with general adaptation to Montana
yield and protein. Protein was negatively correlated with TS,
environment. However, quality characteristics, such as protein
thus protein- or starch-type cultivars may be bred for different
and starch, were not evaluated in that study.
end users. Compared to other cultivars tested, DS Admiral was
Similar to other agricultural commodities, yield and nutri-
the most promising one with above average yield, protein, and RS.
tional characteristics of dry pea affect the benefits of both pro-
ducers and end-users. For farmers, a high-yielding cultivar with
desirable quality for buyers will maximize farm income, while
Core Ideas for end-users various nutritional or chemical characteristics are
Dry pea yield, protein, and resistant starch varied greatly across required for different food and feed purposes. Traditionally,
Montana.
protein has been considered the most important component of
Yield and protein were mainly determined by environments.
pea grains governing end-use quality (Tzitzikas et al., 2006).
Resistant starch is controlled by genetics to a great extent.
In developing countries such as India, for example, dry pea
Effects of drought index, growth period, seed size, and seed
weight on yield, protein, and starch were analyzed. is mainly split for dal as a protein source in the human diet,
whereas in developed countries dry pea is primarily considered
A g ro n o my J o u r n a l Vo l u m e 10 9, I s s u e 4 2 017 1
an excellent high-protein feedstuff (Annicchiarico and knowledge exists regarding the influence of environmental
Iannucci, 2008). In addition to 20% (w/w) protein (on average), factors and agronomic management on the composition of dry
dry pea contains 46% starch and 20% fiber (Tzitzikas et al., pea, especially RS and protein contents.
2006). Although pea is not considered a major source of edible The objective of this study was to investigate the variation of
starch (compared to rice [Oryza sativa L.], wheat [Triticum aes- yield, protein, TS, and RS in dry pea grown in different envi-
tivum L.], corn [Zea mays L.], and potato [Solanum tuberosum ronments across Montana.
L.]), pea starch is now widely processed to noodles in food
industries (Ratnayake et al., 2002). In fact, pea starch is consid- MATERIALS AND METHODS
ered the second best material {after mung bean [Vigna radiata Site Description
(L.) Wilczek]} among all grain legumes for processing starch Field trials were conducted at five experimental loca-
noodles (Tan et al., 2009). tions (Central Agricultural Research Center near Moccasin,
In the process of digestion, a proportion of starch, which Northern Agricultural Research Center near Havre,
is called RS, escapes from digestion in the small intestine of Northwestern Agricultural Research Center near Creston,
healthy individuals within 120 min but fermented in the colon Southern Agricultural Research Center near Huntley, and
(Sun et al., 2015). A literature review conducted by Sajilata et Western Triangle Research Center near Conrad) across the
al. (2006) described that RS has physiological functions simi- state of Montana during 2013 and 2014. Montana is the fourth
lar to those of dietary fibers. Resistant starch can potentially largest state and the largest dry pea producer in the United
help individuals control diabetes and energy balance, and the States (USDA-NASS, 2014) with diverse climate and soil con-
short-chain fatty acids produced by fermenting colonic bacteria ditions. Geographic coordinates and soil types of the experi-
provide direct health benefits to the colon (Birt et al., 2013). mental sites are provided in Table 1.
Due to the considerable health benefits, RS has attracted sub- Weather parameters, including monthly cumulative precipi-
stantial attention in recent years (Haenen et al., 2013; Sun et tation and average temperature during the growing season at
al., 2015). Variable contents of RS among different genotypes each site, are given in Table 2.
of rice (Chiu and Stewart, 2012) and wheat (Hazard et al.,
2012) have been reported. Strydhorst et al. (2015) found winter Experimental Design
pea (cultivar Windham) had lower starch but higher resistant General agronomic practices at the experimental sites are
starch and protein than spring pea cultivars. However, research summarized in Table 3. Starter fertilizer, mainly P, was applied
is needed regarding the variation of RS and protein among dry at Conrad and Creston in 2013 and 2014 based on pre-plant
pea cultivars growing in different environments. Very little soil test. According to previous studies by Chen et al. (2006)
Table 1. Geographic coordinate, altitude and soil type at the experimental locations.
Location Geographic coordinate Altitude Soil type
m
Conrad 4814 N, 11155 W 1117 fine, smectitic, frigid Aridic Argiustoll (Scobey clay loam series)
Creston 4811 N, 11408 W 905 coarse-silty, mixed, Udic Haploboroll (Creston silt loam series)
Havre 4830 N, 10947 W 823 fine-loamy, mixed, superactive, frigid Aridic Argiustoll (Joplin clay loam series)
Huntley 4555 N, 10814 W 920 fine-loamy, mixed, superactive, mesic Aridic Haplustalf (Fort Collins clay loam series)
Moccasin 4703 N, 10957 W 1295 fine-loamy, carbonatic Typic Calciboroll (Judith clay loam series)
Table 2. Monthly precipitation and average temperature at each location in 2013 and 2014 growing season.
Conrad Creston Havre Huntley Moccasin Conrad Creston Havre Huntley Moccasin
Month 2013 2014
Precipitation, mm
Mar. 5 16 0 6 2 20 59 13 55 28
Apr. 9 54 11 30 17 22 19 18 30 16
May 31 84 89 154 80 40 29 12 58 41
June 106 70 142 36 96 63 162 49 71 62
July 28 1 55 16 43 48 12 11 11 34
Aug. 30 23 27 37 24 31 43 96 93 170
Sept. 23 67 38 104 96 50 19 18 19 59
Temperature, C
Mar. 0 1 1 2 0 5 0 3 0 2
Apr. 3 4 4 5 2 4 5 6 8 3
May 10 11 12 14 10 10 11 11 13 9
June 11 14 16 18 14 12 13 14 16 12
July 18 19 20 22 20 19 19 20 22 20
Aug. 19 18 21 23 20 18 18 19 21 18
Sept. 14 14 16 18 15 12 12 13 15 12
treated, and packaged at the coordinator site (Moccasin) then Yij =m + Gi + Ej + lk xik hjk + eij
shipped to the other experimental sites. Seeds were planted k =1
4
General grain Mean grain weight Mean days to flowering
Cultivar Code Seed color size category in this study Maturity in this study Plant height
g per 1000 Table 5. Degree of drought conditions based on
Bridger Br Yellow Medium 212 Mid 64 Medium the modified palmer drought index (PDI).
Cruiser Cr Green Small 197 Mid 64 Tall PDI Wet/dry condition
Daytona Da Green Very large 251 Late 66 Short 4.0 or less Extreme drought
DS Admiral DS Yellow Large 230 Mid 65 Tall 3.0 to 3.9 Severe drought
Jetset Je Yellow Large 226 Late 64 Medium 2.0 to 2.9 Moderate drought
Montech 4152 Mo Yellow MediumLarge 244 Mid 64 Tall 1.9 to +1.9 Near normal
Navarro Na Yellow Very large 255 Early 61 Medium +2.0 to +2.9 Unusual moist spell
Spider Sp Yellow Large 235 Mid 66 Tall +3.0 to +3.9 Very moist spell
SW Midas SW Yellow Medium 202 Mid 65 Medium +4.0 or above Extremely moist
Table 6. AMMI analysis for the effect of environment (E), cultivar (G) and their interaction (G E) on grain yield, resistant starch, protein, and total starch content of peas grown across
Montana in 2013 and 2014. PC1, PC2, PC3, and PC4 are the first, second, third and fourth principal components of the interaction term (G E), respectively.
Source of variation df SOS %SOS P values SOS %SOS P values SOS %SOS P values SOS %SOS P values
E 9 380,794,891 93.1 P < 0.0001 11,188 37.4 P < 0.0001 156,461 92.5 P < 0.0001 170,204 66.9 P < 0.0001
Rep (R) 20 3,086,095 0.4531 205 0.6881 5,043 P < 0.0001 37,066 P < 0.0001
G 8 6,972,736 1.7 P < 0.0001 12,517 41.8 P < 0.0001 2,241 1.3 0.0007 23,049 9.1 P < 0.0001
GE 72 21,000,617 5.1 0.0004 6,205 20.7 P < 0.0001 10,373 6.1 0.0007 61,122 24 P < 0.0002
PC1 16 1,204,8921 57.4 P < 0.0001 2,408 38.8 P < 0.0001 3,912 37.7 0.0001 29,355 48 P < 0.0001
PC2 14 4,009,158 19.1 0.0332 2,107 7 P < 0.0001 2,745 26.5 0.0028 13,021 21.3 0.0114
PC3 12 3,061,822 14.6 0.078 955 3.2 P < 0.0001 1,580 15.2 0.0734 8,187 13.4 0.0992
PC4 10 841,863 4 0.8522 343 1.1 0.0039 906 8.7 0.3217 5,624 9.2 0.2292
Residual 160 24,437,035 2,003 12,453 68,775
Sum of square.
Fig. 1. Additive main effect and multiplicative interaction (AMMI); biplot showing (a) yield, (b) protein, and (c) total starch vs. PC1 of nine
pea cultivars (blue color) and 10 environments (red color). Codes: Bridger (Br), Cruiser (Cr), Daytona (Da), DS Admiral (DS), Jetset (Je),
Montech 4152 (Mo), Navarro (Na), Spider (Sp.), SW Midas (SW). Conrad, 2013 (CO 13), Creston, 2013 (CR 13), Havre, 2013 (HA 13),
Huntley, 2013 (HU 13), Moccasin, 2013 (MO 13), Conrad, 2014 (CO 14), Creston, 2014 (CR 14), Havre, 2014 (HA 14), Huntley, 2014
(HU 14), Moccasin, 2014 (MO 14).
Fig. 2. Relationship between the modified palmer drought index (PDI) and grain yield of pea at various phenological stages ([a]
establishment, [b] vegetative growth, [c] flowering, and [d] grain filling). Regression lines are drawn for each cultivar separately but the
regressions equation represent the case with all data points being considered.
PDI values and degree of drought: 4.0 or less = extreme drought, 3.0 to 3.9 = severe drought, 2.0 to 2.9 = moderate drought, 1.9
to +1.9 = near normal, +2.0 to +2.9 = unusual moist, +3.0 to +3.9 = very moist, and +4.0 or above = extremely moist.
Fig. 3. Relationship (a) between yield and grain weight and (b) between resistant starch (RS) and grain weight of pea. The values are
average number for each cultivar across locations and years.
Protein Content relationship between TS and protein (r = 0.51, P < 0.001) and
The AMMI analysis revealed that 92% of the total varia- between TS and yield (r = 0.15, P = 0.0153).
tion in protein content was related to E, whereas G and GE
explained 1 and 6% of the total variation (P = 0.0007 for DISCUSSION
both), respectively (Table 6). Environment was the most important driving factor in pea
Protein content in Creston (CR13 and CR14) followed by grain yield determination, with a large gap between the average
Huntley (HU13 and HU14) were higher than those in other yield at the high yield potential site and other locations. Hence,
sites (Fig. 1b). Montech 4152 had the greatest content of pro- the genetic and physiological potential of the cultivars are high,
tein while SW Midas showed the lowest protein. Spider had but actual yields were extremely limited by the environmental
greater than average protein content and showed great stability conditions at locations other than Creston. In both years and
for this trait across the environments. during all four distinct phenological stages, Creston had the
There was a significant quadratic relationship between PDI highest PDI values compared to other sites. Relatively moist
and protein at the establishment (Fig. 4a) and at the vegetative condition resulted in a considerably higher yield of all cultivars
growth stages (Fig. 4b). Pearson correlation analysis showed at this site compared to other locations. It should be noted that,
that yield and protein were positively correlated (Table 8). except during the establishment, PDI values did not exceed
1 (even at Creston) in this study, which means that too-wet
Total Starch conditions did not occur in this study. Therefore, drought was
Pea TS was mainly affected by E (67% of the total variation), likely the major cause of huge yield gap between Creston and
nevertheless, the effects of G and GE were also significant at the other locations across Montana. Other environmental
P < 0.01 (Table 6). variables not evaluated in this study, such as soil conditions,
In contrast to RS, range of variation for TS among pea culti- especially soil depth (which affects water holding capacity in
vars was relatively small (Fig. 1c). The highest mean TS content rain-fed systems) may also play important roles in pea yield
was observed at Conrad 2014 (CO14), whereas pea grown in determination. Cultivars responded differently to dry condi-
Creston 2013 (CR13) and Conrad 2013 (CO13) contained tions, suggesting opportunities to select better-adapted culti-
the lowest TS. Several cultivars, such as Cruiser and Jetset, vars and/or avoid planting sensitive cultivars in drier areas.
had large (positive or negative) PC1 values, indicating specific The effect of cultivar on pea grain yield was relatively small,
adaptations (higher TS at certain locations). Averaged over all much smaller than that of environment. On the other hand,
environments, SW Midas had the greatest contents of total the rank of cultivar performance varied greatly by environ-
starch. Pearson correlation analysis (Table 8) showed a negative ment, reflecting site-specific adaptability of cultivars. This sug-
gests that breeding efforts could be more successful if attention
Table 8. Pearson correlation (n = 270) coefficients (and P values is given to the regional interests rather than to make a superior
in italics) between grain yield, protein, resistant starch (RS), and
total starch (TS) in dry pea grown across Montana (nine cultivars, cultivar with general adaptability. For example, in the condi-
2 yr, five experimental sites, and three replications at each site). tion of this study, Navarro outperformed other cultivars in low
yield potential sites (Moccasin, Huntley, and Havre), whereas
Variables Grain yield Protein RS TS
in the highest yield potential environment (Creston), Daytona
Grain yield 1
was the best yielding cultivar. Navarro is among the earliest
flowering pea cultivars in the U.S. NGP with an extended
Protein 0.39 1
flowering period and early maturity (Kandel and Endres,
<0.0001
2010), which likely makes it more resilient to environmental
RS 0.10 0.11 1
constraints. Daytona, on the other hand, has shown excellent
0.0885 0.0776 lodging resistance, which seems to be a crucial prerequisite
TS 0.15 0.51 0.36 1 in high-yielding environments, like Creston. The importance
0.0153 <0.0001 <0.0001 of lodging resistance to achieve a higher yield in pea has been
Fig. 4. Relationship between the modified palmer drought index (PDI) and protein content of pea at the (a) establishment and (b)
vegetative growth stage. Regression lines are drawn for each cultivar separately but the regressions equation represents the case with all
data points being considered. Data of Huntley 2014 were not included. PDI values and degree of drought: 4.0 or less = extreme drought,
3.0 to 3.9 = severe drought, 2.0 to 2.9 = moderate drought, 1.9 to +1.9 = near normal, +2.0 to +2.9 = unusual moist, +3.0 to
+3.9 = very moist, and +4.0 or above = extremely moist.