Field Crops Research 180 (2015) 126–134

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Field Crops Research

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Irrigation frequency and seasonal timing effects on perennial ryegrass

(Lolium perenne L.) seed production
Thomas G. Chastain a,∗ , Craig M. King a , Carol J. Garbacik a , William C. Young III a ,
D.J. Wysocki b
a
Department of Crop and Soil Science, Oregon State University, 107 Crop Science Bldg., Corvallis, OR 97331, USA
b
Department of Crop and Soil Science, Columbia Basin Agricultural Research Center, Oregon State University, PO Box 370, Pendleton, OR 97801, USA

a r t i c l e i n f o a b s t r a c t

Article history: Limited information is available on frequency and seasonal timing of irrigation in perennial ryegrass seed
Received 9 February 2015 crops. Two field experiments were conducted in Oregon to investigate irrigation effects on seed produc-
Received in revised form 22 May 2015 tion in seven cultivars of perennial ryegrass. Three spring irrigation treatments were tested in Experiment
Accepted 26 May 2015
1 over a two-year period: none (control), single irrigation to fill the soil profile to field capacity (FC) at
Available online 15 June 2015
anthesis, and multiple irrigations to maintain soil water deficit ≤50 mm below FC until the beginning of
seed fill; and single fall and spring irrigation timings were tested in Experiment 2 over a three-year period.
Keywords:
Spring irrigation increased seed yield of perennial ryegrass in both experiments while fall irrigation had
Grass seed crop production
Irrigation
no effect in the second experiment. A single spring irrigation increased seed yield by an average 16.1%
Lolium perenne L. whereas multiple irrigations resulted in an average increased yield of 24.6% in Experiment 1. Single spring
Seed yield irrigation in Experiment 2 increased seed yield by an average 6.5%. Seed yield increases attributable to
spring irrigation were the result of a combination of increased seed number and seed weight. Spring
irrigation increased harvest index and seed set. Water use efficiency was reduced by spring and fall
irrigation. Perennial ryegrass seed crops can benefit from spring irrigation but the frequency of irrigation
is best determined by the quantity of water available and economic considerations.
© 2015 Elsevier B.V. All rights reserved.

1. Introduction
Perennial ryegrass is an important cool-season forage and turf
grass seed crop with world-wide annual production of seed on
over 300,000 ha. Traditionally, perennial ryegrass seed crops have
been grown without the aid of irrigation in one of the world’s
principle production regions—Oregon’s Willamette Valley in the
northwestern USA. The Willamette Valley receives 90% of its annual
precipitation (1015 mm) between October and April, with very
little precipitation in late spring and summer during anthesis,
pollination, seed development and fill. The post-harvest period
from August until mid-October is often marked by very dry con-
ditions and there is not enough water available to support optimal
regrowth of the perennial ryegrass for subsequent seed harvests
Abbreviations: HI, harvest index; WUE, water use efficiency; DOY, day of year;
CWU, crop water use; TDR, time domain reflectometry; FC, field capacity; ET, evapo-
transpiration; BBCH, Biologische Bundesanstalt, Bundessortenamt und CHemische
Industrie.
∗ Corresponding author. Tel.: +1 541 737 5730.
E-mail address: Thomas.G.Chastain@oregonstate.edu (T.G. Chastain).
from the stand (Velloza, 1997). Increased ability for seed growers
to apply irrigation water has prompted inquiries about irrigation
recommendations for perennial ryegrass seed crops.
There is limited information available on the effects of irrigation
frequency and seasonal timing on perennial ryegrass seed crop pro-
duction. In England, seed yield of timothy (Phleum pratense L.) and
perennial ryegrass (Lolium perenne L.) was reported to increase with
irrigation (Lambert, 1967; Hebblethwaite, 1977; Hebblethwaite
and McGowan, 1977). Work done recently in New Zealand showed
that first-year perennial ryegrass seed yields are increased with
spring irrigation (Chynoweth et al., 2012). Huettig et al. (2013)
found that spring irrigation strategically timed to coincide with
peak anthesis increased seed yield by as much as 47% in tall fes-
cue [Schedonorus arundinaceus (Schreb.) Dumort.] grown in the
Willamette Valley. Irrigation could be beneficial in fall drought
conditions or when low soil water levels occur during heading,
pollination or seed development in spring.
The typical life of the stand in perennial ryegrass seed crops
vary regionally from one harvest year in the Canterbury Plains of
New Zealand to three harvest years in Oregon’s Willamette Valley.
Tradition, terms of the seed production contract, and other factors
influence the life of the stand in perennial ryegrass seed crops. Work

http://dx.doi.org/10.1016/j.fcr.2015.05.021
0378-4290/© 2015 Elsevier B.V. All rights reserved.
 T.G. Chastain et al. / Field Crops Research 180 (2015) 126–134
done in perennial ryegrass and tall fescue suggests that the seed
yield response of the seed crop to irrigation was influenced only by
the most recent (current crop year) irrigation and not by irrigation
applied in previous harvest years (Hebblethwaite, 1977; Huettig
et al., 2013). In other words, there does not seem to be a carry-over
effect of prior irrigation applications manifested in following years
with regard to seed yield or yield components. Seed yields in cool-
season perennial grass seed crops are generally highest early in the
life of the stand, with declining seed yields as that crop stand ages
(Zapiola et al., 2006).
Present recommendations for irrigation management offer
conflicting crop water use (CWU) requirements for Oregon grass
seed crops. Watts et al. (1968) reported that the CWU requirement
for grass seed production in the Willamette Valley was 281 mm.
Water needs were based on a growing season from March to June.
Cuenca et al. (1992) used a year-round growing season which esti-
mated that CWU required was 762 mm for Willamette Valley grass
seed crops. Unfortunately, both CWU recommendations are based
on water use in grasses managed for turf and not for production
of seed, and are not specific for species such as perennial ryegrass.
Crop water use in tall fescue seed crops was measured and found to
range from 247 mm to 409 mm in a March to June growing season
(Huettig et al., 2013).
The objectives of this study were to (1) determine the effect
of frequency and seasonal timing of irrigation on seed yield and
yield components in seven cultivars of perennial ryegrass, and (2)
develop recommendations for irrigation management in perennial
ryegrass seed production under Oregon conditions.
2. Materials and methods
2.1. Overview
Two field experiments were established at Hyslop Research
Farm near Corvallis, Oregon. Spring irrigation timing and frequency
responses of six cultivars of perennial ryegrass were examined in
Experiment 1 and the seasonal effects of fall and spring irrigation
on a single cultivar of perennial ryegrass were elucidated in Exper-
iment 2. The soil at the site for both experiments was a Woodburn
silt loam (fine-silty, mixed, mesic, Aquultic Argixeroll).
Crop management was based on common production prac-
tices for perennial ryegrass seed production in Oregon’s Willamette
Valley (Young et al., 1999). Development stages of the perennial
ryegrass seed crops in relation to management practices and exper-
imental treatments were characterized by using the BBCH scale
(Hess et al., 1997). Applications of 45 kg N ha−1 (applied as 16-20-
0) were made in October of each crop year on established crops
(BBCH 20-29). Spring nitrogen needs were met by splitting appli-
cations over two dates during March prior to stem elongation (no
later than BBCH 29) in each year for a total N rate of 180 kg N ha−1
(applied as 33-0-0-12). Lodging was controlled with applications of
trinexapac-ethyl plant growth regulator applied at 200 g a.i. ha−1
during early stem elongation (BBCH scale 32-33; Chastain et al.,
2014a).
2.2. Experiment 1
The experimental design for the Experiment 1 was a random-
ized complete-block with a split-plot arrangement of treatments
and four replications. Main plots (18 × 15 m) were three irrigation
treatments and subplots (3 × 15 m) were six perennial ryegrass cul-
tivars. Irrigation treatment main plots were separated by 3-m wide
borders. Experiment 1 was conducted for two consecutive years on
the same plots.
127
Table 1
Timing of irrigation and harvest management events in perennial ryegrass experi-
ments 1 and 2. Events are denoted by the mean day of year (DOY).
Experiment Event Mean DOY
1 Multiple spring irrigations start 137
Multiple spring irrigations end 160
Single spring irrigation application 153–157
Swath plots 187–189
Combine harvest plots 195
2 Single spring irrigation application 140–143
Swath plots 197
Combine harvest plots 210
Single fall irrigation application 260–261
Six perennial ryegrass cultivars (‘Caddieshack’, ‘Cutter’, ‘Derby
Supreme’, ‘Gator 3’, ‘Pirouette’, and ‘SR4500’) were selected in
Experiment 1. Cultivars represented a range of genotypes, plant
morphological characteristics and crop maturity based on date
of anthesis. Crop maturity among cultivars ranged over a two-
week period and ordered from earliest to latest are ‘Cutter’, ‘Derby
Supreme’, ‘SR4500’, ‘Gator 3’, ‘Pirouette’ and ‘Caddieshack’. Plots
were sown with a double-disk drill on 7 May 2002 (DOY 127) at a
seeding rate of 5.6 kg ha−1 in 30-cm rows.
Three irrigation management treatments were employed in
Experiment 1: (i) None, non-irrigated control; (ii) Single, a sin-
gle irrigation application was timed to precede peak anthesis and
filled the soil profile to FC prior to seed filling; and (iii) Multiple,
irrigation water was applied as needed to maintain soil water in
the profile to a deficit ≤50 mm below FC until the start of seed fill.
Irrigation ceased after peak anthesis for both the single and multi-
ple treatments. FC of the soil at the site was 0.46 m3 /m3 (Velloza,
1997). Hebblethwaite and McGowan (1977) had suggested that
50 mm deficits below FC begin to impact transpiration and growth
of perennial ryegrass.
Soil volumetric water content was measured by time domain
reflectometry (TDR) (SoilMoisture Equipment Corp., Santa Barbara,
CA; Musters and Bouten, 2000). Probes for the TDR were installed
horizontally at 15-, 30-, 45- and 60-cm depths in four replica-
tions of ‘SR4500’ and ‘Pirouette’. These cultivars were selected
based on crop maturity (SR4500—early maturity, Pirouette—late
maturity). Measurements were taken weekly, and sometimes more
frequently, throughout the growing season to monitor changes in
soil water. Irrigation treatments were applied with a Pierce Corpo-
ration Acremaster MicroLinear (Eugene, Oregon). The MicroLinear
had three spans with individual valves allowing control over each
span. Each span had six D3000 spray heads (Nelson Irrigation Cor-
poration, Walla Walla, Washington); two at the end of the span
were half circle to prevent water application between main plots.
Pressure regulators (6.9 × 104 Pa) and nozzles limited water output.
Initially in Year 1 output was 11.9 L min−1 and output was increased
to 24.1 L min−1 by installing new nozzles. Application rates were
further controlled by altering MicroLinear speed.
Expected application rates were determined by using engineer-
ing specifications for sprinkler output and rate of linear movement.
In addition, field measurements were collected by randomly pla-
cing containers along the span of the linear and measuring collected
water. Values for water collected in containers were within 10% of
those calculated from sprinkler specifications. Soil water content
was used to schedule spring irrigation (Table 1).
2.3. Experiment 2
The experimental design for Experiment 2 was a randomized
complete-block with four replications. A pre-plant application
of fertilizer (16-20-0) was made at a rate of 224 kg ha−1 dur-
ing seedbed preparation. ‘Evening Shade’ perennial ryegrass was
128 T.G. Chastain et al. / Field Crops Research 180 (2015) 126–134

planted at 5.6 kg ha−1 in 30-cm spaced rows on 30 September 2008
(DOY 274). Activated charcoal was applied in a 3-cm wide band over
the seeded rows at the time of planting at a 28 kg ha−1 rate. The pur-
pose of the charcoal was to protect the emerging grass seedlings
from diuron (N -[3,4-dichlorophenyl]-N, N-dimethylurea), which
was applied at a 2.24 kg ha−1 rate on 1 October 2008 (DOY 275)
to control weeds during establishment of the crop. Experiment 2
was conducted for three consecutive years on the same plots.
Irrigation treatments in Experiment 2 were applied in different
seasons: (i) None, non-irrigated control; (ii) spring, a single spring
irrigation application timed to precede peak anthesis and filled the
soil profile to FC prior to seed filling; and (iii) fall, irrigation dur-
ing September and October to alleviate precipitation deficits. Each
irrigation treatment plot was 3 × 15 m. Irrigation applications were
made in the same manner as in Experiment 1.
2.4. Seed yield components
Two samples were cut at ground level from each plot just prior
to anthesis to determine fertile tiller number and above-ground
biomass in every year of Experiment 1 and Experiment 2. Two crop
rows were hand harvested in 30-cm2 quadrats and oven-dried at
65 ◦ C for 48 h. Fertile tillers were separated from vegetative tillers,
counted and weighed.
Samples were also collected for analysis of spikes prior to anthe-
sis in Experiment 1. Approximately 15 spikes were selected at
random from each plot. Spikes were frozen at −15 ◦ C until anal-
ysis. Ten spikes were used to determine the number of spikelets
spike−1 . Two spikelets were selected at the top, middle and bottom
of four panicles to determine the number of florets spikelet−1 .
divided by CWU (estimated) for each of the seven cultivars and
irrigation management treatments.
2.7. Statistical analysis
Analysis of variance was conducted in Experiment 1 and 2 by
using PROC GLM from SAS (SAS Institute, 2009). With the exception
of interactions, treatment means were separated by Fisher’s pro-
tected LSD values at the 5% level of significance. Stability analysis
was conducted to assess the seed yield stability of the perennial rye-
grass cultivars across irrigation environments (year and irrigation
treatment combinations) and interactions according to the method
of Eberhart and Russell (1966). Mean values of seed yields for each
cultivar were regressed on environmental means, the average seed
yield recorded across cultivars for each year and irrigation treat-
ment combination. Finally, the regression coefficients (slope) were
used in determining the stability index for perennial ryegrass cul-
tivars, and plotted against mean seed yields across irrigated and
non-irrigated environments.
3. Results
3.1. Climatic conditions
Spring weather during the two experiments was characterized
by mostly wetter than normal conditions (Fig. 1). In each spring in
Experiment 1, ET varied less early in the season when conditions

2.5. Seed yield and harvest index

Plots were harvested by swathing when seed moisture content

reached approximately 400 g kg−1 . A commercial swather, modi-
fied for small plots, was used to harvest plots into windrows for
drying. Two dates were used in Experiment 1 to accommodate
the difference between the earlier maturing ‘Cutter’ and ‘Derby
Supreme’ cultivars and the remaining later-maturing cultivars. The
average swathing dates for each experiment are shown in Table 1.
A plot combine was used to thresh seed when seed moisture
in the swath had decreased to 120 g kg−1 . Harvested yield was
measured directly in the field with a sub-sample being collected
for further analysis in the laboratory. Sub-samples were cleaned
to marketable seed yield quality standards by using a laboratory
size Clipper M-2B (A.T. Farrell, Saginaw, MI). Cleanout from the
conditioning process was used to calculate clean yield. Samples
to determine seed weight were hand cleaned using screens and
a blower prior to counting. The seeds in two 1000 samples from
each plot were counted by an electric seed counter (The Old Mill
Company, Savage, MD) and weighed, and seed weight was calcu-
lated. Harvest index was calculated as the ratio of clean seed yield
to above-ground biomass. Samples were cleaned to remove soil and
dead plant tissue prior to drying at 65 ◦ C for 48 h before weighing.

2.6. Water use efficiency

The crop growing season was defined to start on 15 March (DOY

75) and end on the day of swathing. Drainage and runoff are negligi-
ble for the soil type and lack of slope, and were not considered in the
calculation of CWU. The calculation also does not account for water
used prior to TDR measurements when the soil was saturated.
Evapotranspiration (ET) reported by Agrimet (2013) at Hyslop Farm
Fig. 1. Cumulative precipitation at Hyslop Farm during the spring (top) and fall
was used to estimate this water use. From 15 March (DOY 75)
(bottom) irrigation period. The mean refers to the long-term average for the period.
until the first TDR measurement, ET was summed and then added Experiment 1 was conducted in Year 1 and Year 2. Experiment 2 was conducted in
to CWU. Water use efficiency was calculated as clean seed yield Year 3, 4, and 5.
 T.G. Chastain et al. / Field Crops Research 180 (2015) 126–134
Fig. 2. Spring irrigation effects on soil water content in Year 1 (Experiment 1) at 15-, 30-, 45-, and 60-cm depths in the soil profile under no irrigation (none), a single
strategically timed irrigation (single), and multiple irrigations through the period (multiple). Soil water values are averaged across cultivars ‘SR4500’ and ‘Pirouette’.
are typically wetter and cooler in the Willamette Valley (data not
shown). The magnitude of ET variation was much greater late in the
season in the period immediately prior to crop harvest when con-
ditions are typically warmer and drier. In Year 1, the spring season
was wet in the first half (April rainfall, DOY 91–120, was 216% of
normal), and dry during the second half. May (DOY 121–151) and
June (DOY 152–181) rainfall were 54 and 21% of normal, respec-
tively. This corresponded with reduced soil water beginning near
the onset of anthesis (DOY 145) (Fig. 2).
In Experiment 1, Year 2 was characterized by alternating wet
and dry periods throughout, but overall lower than normal rainfall
and as a result these conditions caused soil drying earlier in the
season. Rainfall in March (DOY 61–91) and May (DOY 122–152)
was low, averaging 45 and 68% of normal, respectively. Precipi-
tation in April (DOY 92–121) was near normal (94%) while June
(DOY 153–182) precipitation was above normal (122%). Soil water
declined earlier in Year 2 and was being drawn down during stem
elongation (DOY 110) through seed fill (DOY 185) in non-irrigated
plots (Fig. 3).
Rainfall exceeded the long-term mean for the area during the
spring (April–June) period in each of the three years of Experiment
2. April–June rainfall was 179, 151, and 163% of normal in Year
3, Year 4, and Year 5, respectively. Late summer and early fall is
often a relatively dry season in Oregon, but during Experiment 2,
the conditions were drier than normal (Fig. 1).
3.2. Experiment 1—Spring irrigation and cultivars
The wet conditions experienced in Year 1 (Fig. 1) made early
season irrigations unnecessary, so the single application was made
between DOY 155 and DOY 158 and the multiple applications were
made between DOY 154 and DOY 163 (Table 1). Irrigation had a
129
direct and proportional effect on the soil water throughout the pro-
file (Fig. 2). Dry conditions in Year 2 were conducive to both single
and multiple irrigation treatments (Fig. 3). The single irrigation in
Year 2 was applied between DOY 152 and DOY 156. Multiple irri-
gations began just prior to the start of pollination on DOY 120 and
ended on DOY 156. The average water applied in single strategic
irrigation was 94 mm whereas the average of multiple irrigations
was 166 mm.
Soil water content of single-irrigated plots in Year 1 ranged from
34.9% at 15-cm depth to 38.6% at 60-cm depth on DOY 157 (peak
anthesis) (Fig. 2). By comparison, for plots receiving no irrigation,
soil water content ranged from 16.6% at 15-cm depth to 31.7% at
60-cm depth at peak anthesis. By the final date of swathing on DOY
189 in Year 1, soil water content had declined to 11.0 and 14.0%,
for none and single treatments, respectively, at the 15-cm depth.
Also at that date, soil water dropped to 24.4 and 25.3% for none and
single treatments, respectively, at the 60-cm depth (Fig. 2).
In neither year did plants have visible signs of wilting or stress,
though leaf senescence began earlier in non-irrigated plants than in
irrigated plants. Despite differences in weather between years, soil
water content had declined to similar levels in the single irrigation
treatment by the time of water application on DOY 154 in Year
1 and on DOY 152 in Year 2, respectively (Fig. 3). Average peak
anthesis for the crop was the same in Year 2 as in Year 1 (DOY 158).
There were no differences in soil water depletion or crop maturity
between ‘SR4500’ and ‘Pirouette’ as measured by TDR so the soil
water content values presented in Figs. 2 and 3 are the average of
the two cultivars.
Significant year effects were noted for seed yield, seed weight,
seed set, HI, and WUE, but several of these effects including
seed yield, seed weight, HI and WUE were best explained by
Year × Irrigation or Year × Cultivar interactions (Table 2). Seed set
130 T.G. Chastain et al. / Field Crops Research 180 (2015) 126–134

Fig. 3. Spring irrigation effects on soil water content in Year 2 (Experiment 1) at 15-, 30-, 45-, and 60-cm depths in the soil profile under no irrigation (none), a single
strategically timed irrigation (single), and multiple irrigations through the period (multiple). Soil water values are averaged across cultivars ‘SR4500’ and ‘Pirouette’.

was increased from 19.7% in Year 1 to 29.8% in Year 2 (data not
shown). The increased seed set in Year 2 may have been the result
of better pollination conditions in Year 2 because of lower rainfall
during anthesis.
Irrigation had no effects on several seed yield components
including spikes m−2 , spikelets spike−1 and florets spikelet−1
(Table 2). Irrigation treatments were applied after spike emergence
(BBCH 50) and thus, could not have been influential in determin-
ing the number of spikes as well as spikelets spike−1 or florets
spikelet−1 as these seed yield components were already set prior
to irrigation. While irrigation treatments altered the canopy struc-
ture by increasing vegetative matter, possibly leading to reduced
air movement through the stand, the incidence and severity of crop
diseases and other pests were not observed to be influenced by
irrigation treatment.
Averaged over the two-year period in Experiment 1, seed yield
was increased by both single and multiple irrigations with the best
results observed from multiple irrigations (Table 3). A single strate-
gic irrigation timed to fill the soil profile to FC at anthesis increased
seed yield by an average of 16.1% over the non-irrigated control
while multiple irrigations resulted in yield increases that averaged
24.6%. Seed yield was different among years (Table 2) with yields
in Year 1 being an average 217 kg ha−1 greater than in Year 2 (data
not shown).

Table 2
Analysis of variance for irrigation, cultivar, and seasonal irrigation timing (season) effects on seed yield and other seed production characteristics in perennial ryegrass.
Experiment 1 was conducted in Years 1 and 2, and Experiment 2 was conducted in Years 3, 4, and 5.

Source of variation Seed yield Spikes Spikelets spike− Florets spikelet− Seed set Seed number Seed weight HI WUE

Experiment 1
* ** ** *** **
Year (Y) ns ns ns ns
*** ** ** *** * ***
Irrigation (I) ns ns ns
Y×I ns ns ns ns ns ns ns * **
*** *** ** ** *** *** *** *** ***
Cultivar (C)
Y×C ***
ns ns ns ns *** ** * ***

C×I ns ns ns ns ns ns ns ns ns
Y×C×I ns ns ns ns ns ns ns ns ns

Experiment 2
* * *
Year (Y) ns – – – ns ns
* * * * **
Season (S) ns – – –
Y×S ns ns – – – ns ns ns ns

ns = not significant.
*
P ≤ 0.05.
**
P ≤ 0.01.
***
P ≤ 0.001.
 T.G. Chastain et al. / Field Crops Research 180 (2015) 126–134 131

Table 3
Effects of spring irrigation (none, single, and multiple) on seed yield and seed pro-
duction characteristics in six cultivars of perennial ryegrass in Experiment 1. Means
of seed production characteristics are averaged over two years.

Irrigation

Characteristic None Single Multiple

−1
Seed yield (kg ha ) 1639a† 1903b 2042c
Seed number (×104 m−2 ) 10.5a 11.5b 11.9b
Seed weight (mg) 1.63a 1.73b 1.78c
Seed set (%) 21.9a 26.2b 26.1b
Spikes m−2 2631a 2627a 2502a
HI 12.2a 14.1b 13.5b
WUE (kg ha−1 mm−1 ) 5.82c 5.30b 4.78a
†
Means within seed characteristics followed by the same letter are not significantly
different by Fisher’s LSD values (P = 0.05).

Spring irrigation increased seed number m−2 over the control

in perennial ryegrass but there were no differences among single
and multiple irrigations (Table 3). Seed weight was incrementally
increased with single and multiple irrigations with the greatest
seed weight observed with multiple irrigations (Table 3). This
increase in seed yield as a result of spring irrigation was likely due to
an 11.4% increase in seed number, and increases in seed weight that
ranged from 6.1 to 9.2%. The increased number of seed produced
was possibly attributable to a 19.4% increase in seed set as a result
of spring irrigation. No differences in seed set were found between
single and multiple irrigations. Partitioning to seed was increased
by irrigation as HI was improved by 13.1% with spring irrigation.
Incremental reductions in WUE were evident with irrigation and
the greatest reduction in WUE was found with multiple irrigations
(Table 3). Year × Irrigation interactions were significant for HI and Fig. 4. Regression of mean cultivar seed yield on irrigation environments (top) and
WUE (Table 2). These were largely the result of greater absolute classification of perennial ryegrass cultivars based on mean seed yield and stability
index (bottom) in Experiment 1. Environmental means are the mean seed yields of
range in differences among irrigation treatments for HI in Year 2 the cultivars for each year and spring irrigation treatment combination. The fitted
(23.0%) than in Year 1 (6.9%), and for WUE in Year 2 (29.7%) than in regression line for ‘Derby Supreme’ is dashed for comparison purposes in the top
Year 1 (6.5%). part of the figure. The dashed lines in the bottom part of figure represent the overall
Averaged over the two years in Experiment 1, seed yield var- mean seed yield (vertical) and stability index = 1.0 (horizontal).
ied among cultivars with Pirouette having the highest yields while
Derby Supreme, Gator 3 and SR 4500 had among the lowest yields
(Table 4). Pirouette’s low seed weight was offset by other char- sensitivity to irrigation environment than the other cultivars. To
acteristics that contributed to high observed seed yield including further examine the seed yield responses of cultivars in Experiment
high seed number m−2 , seed set, and spike production. Caddieshack 1 and in particular the Cultivar × Year interaction for seed yield,
also had high seed yields but had higher seed weight than Pirou- the mean seed yield of irrigated and non-irrigated environments
ette. Characteristics that likely contributed to high seed yield in was plotted against the stability index derived using the method-
Caddieshack included spike production, seed weight, seed number ology of Eberhart and Russell (1966). Application of the stability
m−2 and seed set. Both cultivars had high HI while a low yielding index in the classification of the cultivars reveals that ‘Pirouette’ and
cultivar, Gator 3 also had high HI. Among cultivars, there was some ‘Caddieshack’ had the greatest seed yield stability with regard to
variation in WUE with the highest yielding cultivars making the yield performance across irrigated and non-irrigated environments
best use of the water available. (Fig. 4). ‘Derby Supreme’ had not only the lowest and most variable
The regression of seed yield of cultivars on irrigation environ- seed yields but also the lowest seed yield stability across environ-
ment (combinations of year and presence or absence of spring ments. ‘Gator 3’ had the least seed yield stability among the other
irrigation) revealed that five of the six cultivars tested demon- five cultivars tested but was clearly superior to ‘Derby Supreme’
strated a strong and positive yield response to the application of in that regard. Seed yield performance under irrigated manage-
spring irrigation (Fig. 4). The cultivar ‘Derby Supreme’ had less ment is not an objective of perennial ryegrass plant improvement

Table 4
Cultivar effects on seed yield and seed production characteristics in perennial ryegrass in Experiment 1. Means of seed production characteristics are averaged over two
years.

Cultivar Seed yield (kg ha−1 ) Spikes (m−2 ) Seed weight (mg) Seed number (m−2 ) Seed set (%) HI WUE (kg ha− 1 mm−1 )
No No × 104

Cutter 1852b† 2485ab 1.88d 10.2b 21.8ab 12.1ab 5.28b

Pirouette 2026c 2649b 1.59b 13.1e 27.5de 14.7c 5.77c
Derby Supreme 1754a 2379a 1.86d 9.8a 20.2a 11.4a 5.00a
SR 4500 1783ab 2396a 1.70c 10.8c 28.7e 12.8b 5.07a
Caddieshack 1975c 2638b 1.72c 11.9d 25.7cd 14.7c 5.61c
Gator 3 1779a 2973c 1.52a 12.1d 24.5bc 13.6bc 5.06a
†
Means within columns followed by the same letter are not significantly different by Fisher’s LSD values (P = 0.05).
132 T.G. Chastain et al. / Field Crops Research 180 (2015) 126–134
Table 5
Seasonal irrigation effects (none, fall, and spring) on seed yield and seed production
characteristics in one cultivar of perennial ryegrass in Experiment 2. Means of seed
production characteristics are averaged over three years.
Irrigation
Characteristic None Fall
−1 †
Seed yield (kg ha ) 1620a 1591a
Seed number (×104 m−2 ) 9.0a 8.7a
Seed weight (mg) 1.80a 1.83ab
Spikes m−2 2627a 3028a
HI 13.5ab 11.9a
WUE (kg ha−1 mm−1 ) 5.62b 4.32a
†
Means within seed characteristics followed by the same letter are not significantly
different by Fisher’s LSD values (P = 0.05).
programs but it is interesting that there is this much variation
among cultivars in sensitivity to irrigation management.
3.3. Experiment 2—Seasonal irrigation timing
The single spring irrigation application averaged 102 mm while
fall irrigation applications averaged 127 mm in Experiment 2. There
were significant effects of Year on the spike production, HI, and
WUE in Experiment 2 (Table 2), but there were no significant
Year × Season (none, fall, spring irrigation treatment) interactions
on any characteristic measured.
Averaged over the three-year period of Experiment 2, a single
strategic spring irrigation increased seed yield by 6.5% over the non-
irrigated control (Table 5). The increase in seed yield as a result of a
single spring irrigation was much less than observed in Experiment
1. A possible reason for this difference was that spring rainfall in
Experiment 2 exceeded 150% of the long-term average in all three
years of the trial (Fig. 1). Like in Experiment 1, seed number m−2 and
seed weight appeared to be the primary contributors to the increase
in seed yield as a result of the single spring irrigation (Table 5). Seed
number was increased by 4.4% and seed weight was increased by
2.8% with the single spring irrigation. Spikes m−2 were not affected
by the single spring irrigation nor was HI. Spring irrigation caused
a 21% decrease in WUE.
Fall irrigation had no effect on perennial ryegrass seed yield or
on other seed production characteristics in Experiment 2 (Table 5).
The lack of response to fall irrigation was observed despite much
drier than normal conditions in every fall during Experiment 2
(Fig. 1). Fall irrigation produced seed weight that was not different
from spring irrigation or the no-irrigation control (Table 5). Like
spring irrigation, fall irrigation caused a large decline in WUE but
was not offset by an increase in seed yield.
4. Discussion
4.1. Irrigation and soil water depletion
Soil water depletion was not as great for perennial ryegrass in
this study as was observed under similar conditions in tall fes-
cue seed crops (Huettig et al., 2013). Garwood and Sinclair (1979)
observed soil water depletion by perennial ryegrass in a well-
draining sandy loam soil to a depth of 80 cm, but when grown as a
seed crop in the seasonally saturated soils of the Willamette Val-
ley, more than 90% of perennial ryegrass root biomass was found
to be located in the top 30-cm of the soil profile (Garbacik et al.,
2003). The more rapid rate of soil water content decline in single
irrigated plots than in non-irrigated plots suggested some deple-
tion of the applied water by the crop. Depletion of soil water by the
crop might have taken place below the 60-cm depth monitored by
TDR in this study, though the amount was probably negligible as
few roots likely extended to that depth. As a result of high rainfall
in April during Year 1, high residual levels of water were retained
in the soil well into the month of May despite the dry conditions
in that period and so soil water depletion was not high in that
year.
Spring
4.2. Seed yield and harvest index
1726b
9.4b
1.85b Perennial ryegrass seed yield was consistently increased by
2629a spring irrigation despite seasonal rainfall (April–June) that was con-
14.5b siderably greater than the long-term average for the location in
4.46a
four of five years (Fig. 1). Multiple irrigations in spring proved to be
more beneficial in improving perennial ryegrass seed yield than a
single irrigation timed to support flowering and seed yield. A sin-
gle strategic irrigation to support the critical phases of reproductive
development increased seed yield in tall fescue but multiple irriga-
tions did not provide additional yield benefit (Huettig et al., 2013).
While the single irrigation produced lower seed yields in perennial
ryegrass than multiple irrigations, this lower level of water appli-
cation might be an attractive option for seed growers that have
limited water availability. Economic considerations might also play
a role in choosing to apply a lower water quantity with the single
irrigation yet still producing increased seed yield.
Chynoweth et al. (2012) examined spring irrigation in peren-
nial ryegrass seed production on a light-textured soil in New
Zealand’s Canterbury Plains. Spring irrigation totaling 205 mm (66%
ET replacement) produced a 26.1% increase in perennial ryegrass
seed yield that was comparable to 24.6% yield increased observed
in this study with 166 mm applied in the multiple irrigation treat-
ment. While seed yield increase was similar among the studies, the
difference in the applied water required to achieve this increase
was likely attributable to the heavier-textured silt loam soil and
greater water holding capacity than the Canterbury Plain soil.
Hebblethwaite (1977) demonstrated that perennial ryegrass
seed crop yield responses to irrigation varied among years based on
growing season precipitation. Differences in temperature, amount
and timing of precipitation and varying disease pressures between
years can impact the beginning of spring growth, yield component
development and pollination.
Irrigation treatments coincided with anthesis and the beginning
of seed fill, reducing the potential for water deficit (Figs. 2 and 3)
during the seed fill process. The timing of water deficit stress in
perennial ryegrass determines the effect on seed yield components
(Hebblethwaite, 1977; Rowarth et al., 1997). Hebblethwaite (1977)
found that water stress at multiple times during crop development
were additive in decreasing seed yield, but certain stages were more
sensitive than others.
Fall irrigation had no influence on seed yield despite dry con-
ditions in all three years tested (Fig. 1). Perennial ryegrass has
been shown to produce greater quantities of carbohydrates for
fall regrowth than tall fescue under drought conditions, causing
poorer growth in the following spring (Karsten and MacAdam,
2001). Irrigation during the fall regrowth period is thought to alle-
viate some of this stress and has been shown to improve tillering
as well as other measures of regrowth in perennial ryegrass seed
crops (Velloza, 1997). Nevertheless, these benefits of fall irrigation
in perennial ryegrass were not materialized in greater seed yields
in the following harvest season.
Harvest index is a measure of grass seed crop management
impacts on dry matter partitioning to seed in relation to total above-
ground biomass (Zapiola et al., 2006). Spring irrigation increased
HI in both experiments. Hebblethwaite (1977) found that spring
irrigation in perennial ryegrass seed crops increased HI from 7%
to 11%. No responses of HI to spring irrigation were noted by
Chynoweth et al. (2012) in perennial ryegrass seed crops and by
Huettig et al. (2013) in tall fescue seed crops.
 T.G. Chastain et al. / Field Crops Research 180 (2015) 126–134
Fig. 5. Relationship of seed number m−2 and seed yield in seven cultivars of peren-
nial ryegrass over a five-year period in Experiments 1 and 2. Values are none = no
irrigation, spring = spring irrigation, and fall = fall irrigation.
4.3. Components of seed yield
Management practices for seed production of perennial ryegrass
and other grasses that take place early in crop development can
affect production of fertile tillers m−2 , spikelets m−2 , and florets
m−2 (Chastain and Grabe, 1989; Young et al., 1999; Chastain et al.,
2011), while those timed late in reproductive development such as
spring irrigation and PGR applications typically do not affect fertile
tillers m−2 or spikelets m−2 (Huettig et al., 2013; Chastain et al.,
2014a,b).
Seed number was the most important component that led
to increased perennial ryegrass seed yield in response to spring
irrigation in this study (Fig. 5). Spring irrigation increased seed
number and seed weight in tall fescue, and contributed to higher
seed yields (Huettig et al., 2013). The importance of seed number
as a contributor to seed yield in perennial ryegrass has been pre-
viously demonstrated for other management practices but not for
irrigation (Young et al., 1999; Chastain et al., 2014a, 2014b). Despite
the differences observed among cultivars and years for irrigation
management, the relationship between seed yield in perennial rye-
grass and seed number is evident.
Seed weight also made contributions to seed yield increases in
perennial ryegrass as a result of spring irrigation. Lambert (1967)
found that irrigation increased the proportion of heavy seed in
timothy (Phleum pretense L.) and Chynoweth et al. (2012) dem-
onstrated that seed yield increases resulting from irrigation in
perennial ryegrass were associated with increases in seed weight.
Rowarth et al. (1997) did not observe differences in seed weight
from seed harvested from water-stressed and non-stressed peren-
nial ryegrass plants grown under greenhouse conditions.
Assimilates are not considered to be limiting in seed fill pro-
cesses of perennial ryegrass leading to increased seed weight
(Warringa et al., 1998), as the inflorescence is the primary pho-
tosynthetic organ during this period. Inflorescence assimilates are
thought to be supported by mobilization of pre-anthesis assimi-
lates and current assimilates from leaves and stems during stress
(Clemence and Hebblethwaite, 1984). However, partitioning to
seed in perennial ryegrass and consequent seed filling is not
dependent on the mobilization of water soluble carbohydrates from
the internodes to developing seed, and in fact, storage in the spike
is sufficient to fill all available seeds (Trethewey and Rolston, 2009).
Increased seed weight has been attributed to longer seed fill
periods. Warringa et al. (1998) found that differences in seed weight
within a perennial ryegrass spikelet could be partially attributed to
different flowering times. Later senescence of the irrigated stands of
perennial ryegrass may have led to increased seed weight as a result
133
of a longer seed fill period. Delayed harvest has been reported in
irrigated stands of timothy (Lambert, 1967) and perennial ryegrass
(Hebblethwaite, 1977).
4.4. Water use efficiency
Crop water use averaged 267 mm with no irrigation and aver-
aged 363 mm with a single spring irrigation treatment. These CWU
values were based on the assumptions that the effective rooting
depth for perennial ryegrass managed for seed production under
conditions of the Willamette Valley was 60 cm. Perennial ryegrass
seed crop CWU values were similar to those measured for tall fes-
cue seed crops under similar conditions (Huettig et al., 2013) and
not different from values extrapolated from grasses managed for
turf in the region (Watts et al., 1968).
Irrigation reduced WUE in perennial ryegrass seed crops in both
experiments. In lysimeter studies, WUE for perennial ryegrass seed
production was found to vary with rate of applied N with higher
rates producing greater WUE than low rates (Cookson et al., 1997).
Interaction of spring irrigation treatment and cultivar influenced
WUE in tall fescue seed crops (Huettig et al., 2013). Values for
WUE ranged from 3.3 to 9.9 kg ha−1 mm−1 among spring irrigation
and cultivars in tall fescue, and the effects were mixed-irrigation
increased, decreased or had no effect on WUE. Perennial ryegrass
WUE values were found to be in a similar range in this study to
those in tall fescue and are within the same range as WUE reported
for wheat: 2.5 to 18.3 kg ha−1 mm−1 (Hatfield et al., 2001).
5. Conclusions
This study demonstrates that spring irrigation increases seed
yield of perennial ryegrass under Oregon conditions. Six of the
seven cultivars tested responded to spring irrigation with increased
seed yield. The magnitude of the seed yield response to spring
irrigation was dependent on growing season precipitation; most
importantly, precipitation near anthesis and early seed fill. Avail-
able water during these developmental stages increased seed yield
and served as indicators of the sensitivity of these periods to water
deficits. Nevertheless, seed yield increases were noted with spring
irrigation even under conditions that were wetter than normal for
the region. On the other hand, fall irrigation did not improve seed
yield despite very dry fall weather conditions.
Seed number and seed weight in perennial ryegrass were
influenced positively by spring irrigation, and contributed to the
observed seed yield increases in this study. Irrigation and other
management practices should aim to increase harvestable seed
number. This information will serve as the basis for recommen-
dations on irrigation management in perennial ryegrass seed crops
grown in Oregon’s Willamette Valley.
Acknowledgements
This work was funded in part by grants from the George
R. Hyslop Professorship for Oregon Grass Seed Research and
Education—OSU Foundation, Oregon State University, and the Ore-
gon Seed Council.
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