J Agro Crop Sci (2016) ISSN 0931-2250

DROUGHT STRESS

Morphological and Physiological Responses of Plantain

(Plantago lanceolata) and Chicory (Cichorium intybus) to
Water Stress and Defoliation Frequency
L. M. Cranston1, P. R. Kenyon1, S. T. Morris1, N. Lopez-Villalobos1 & P. D. Kemp1,2
1 Sheep Research Centre, Institute of Veterinary, Animal and Biomedical Sciences, Massey University, Palmerston North, New Zealand
2 Institute of Agriculture and Environment, Massey University, Palmerston North, New Zealand

Keywords Abstract
drought tolerance; evapotranspiration rate;
photosynthetic rate; relative growth rate; Plants are often subjected to periods of water stress. There are little data examin-
taproot diameter ing the effect of water stress on the forage species Plantago lanceolata and Cichori-
um intybus. In two pot experiments with P. lanceolata and C. intybus,
Correspondence morphological responses under optimum, dry, and very-dry water treatments
L. M. Cranston
with weekly, fortnightly and 3-weekly defoliation intervals and physiological
Sheep Research Centre
responses under optimum and very-dry water treatments were measured. A third
Institute of Veterinary
Animal and Biomedical Sciences experiment compared the rooting depths of P. lanceolata and C. intybus under
Massey University field conditions. These findings suggest that both P. lanceolata and C. intybus
Private Bag 11-222 can survive and continue to grow under water stress conditions with the main
Palmerston North 4442 differences between the two species being attributable to morphological charac-
New Zealand teristics (root mass, taproot diameter and shoot mass fraction) rather than differ-
Tel.: +64 6 951 85809
ences at a physiological level. Overall, the results suggest plantain may be more
Email: l.cranston@massey.ac.nz
productive under moderate drought due to its greater shoot mass fraction,
Accepted March 2, 2015 whereas chicory may be more productive and persistent under severe drought
due to its greater root mass, taproot diameter and root depth under field condi-
doi:10.1111/jac.12129 tions.

greater understanding of how plants respond to moisture

Introduction
Plants are often subjected to periods of soil and atmo-
spheric water deficits with drought posing a significant
environmental constraint to plant survival and crop pro-
ductivity (Chaves et al. 2003). This will likely become
increasingly important as the world is challenged by pre-
dicted warmer climates (IPCC 2007). Temperate pastoral
systems in New Zealand are predominantly based on
perennial ryegrass (Lolium perenne L.) with a relatively
minor component of white clover (Trifolium repens L.)
(Kemp et al. 2002). However, throughout summer and
autumn production of ryegrass/white clover pasture can be
limited in both quality and quantity, resulting in reduced
feed intake and poor animal performance (Hughes et al.
1980, Burke et al. 2002, Moorhead et al. 2002).
The occurrence of morphological and physiological
responses, which may lead to some adaptation to
drought stress, can vary considerably among species. A
stress could play a major role in stabilising pasture per-
formance under drought conditions. Forage chicory
(Cichorium intybus L.) and plantain (Plantago lanceolata
L.) are widely used throughout the world as high feed
quality perennial herbages (Sanderson et al. 2003, Lab-
reveux et al. 2006, Li et al. 2010, Golding et al. 2011,
Hutton et al. 2011). Both plantain and chicory are tap-
rooted (Stewart and Charlton 2006). The health and car-
bohydrate storage of taproots influence the persistence of
plants (Lodge 1991) and also confer a degree of drought
tolerance (Nie et al. 2008) through accessing water dee-
per in the soil profile. Consequently, both chicory and
plantain are relatively tolerant to moisture stress (Mook
et al. 1989, Nie et al. 2008). However, Nie et al. (2008)
reported that chicory can tolerate moisture stress to a
greater degree than plantain.
However, overall little is known about effects of
water stress and defoliation frequency and the potential

© 2015 Blackwell Verlag GmbH, 202 (2016) 13–24 13

Cranston et al.
interaction between these on the morphological and
physiological responses of both plantain and chicory.
Therefore, the objective of this research was to address
these issues by studying effects of water stress and defoli-
ation frequency on the herbage production, biomass allo-
cation, persistence and physiological responses of chicory
and plantain under glasshouse conditions, and to assess
the rooting structure of chicory and plantain under field
conditions.
Materials and Methods
Two glasshouse experiments were conducted in the Plant
Growth Unit, Massey University, Palmerston North, New
Zealand (latitude 40°230 S). In addition, one field experi-
ment was carried out at the Massey University Pasture and
Crop Research Unit, 5 km south of Palmerston North,
New Zealand.
Experiment 1
Plant establishment
The experiment was conducted between 24 February and
22 August 2012. Two plant species, plantain cv. ‘Ceres
Tonic’ and chicory cv. ‘Puna II’, were used. The potting
medium consisted of 33 % Manawatu silt loam (B Hori-
zon), 33 % coarse sand (3 mm) and 33 % fine sand
(<0.5 mm) amended with 100 g short term fertiliser mix
(Woodace, flowering plant special; LebanonTurf; 14 % N,
6 % P, 11.6 % K + trace elements) and 50 g dolomite for
30 l of potting medium as a base start fertiliser. Prior to
seed establishment (17 February 2012), soil samples of the
potting medium were taken to determine soil water reten-
tion and pF curve. Field capacity was reached at a gravimet-
ric water content of 11 g per 100 g.
The pots were 250 mm in diameter and 250 mm in
depth and were filled with 10.0  0.05 kg of potting med-
ium. Seeds were sown directly into 154 pots (76 chicory
and 76 plantain), on 24 February 2012, and pots were hand
watered daily with a sprinkler. After seeds germinated,
seedlings were thinned to six per pot and then to four per
pot when well established on 22 March 2012 (resulting in a
total of 304 chicory plants and 304 plantain plants).
An automatic watering system incorporating an individ-
ual dripper in each pot was established on 5 March 2012.
All pots were regularly watered to maintain 100 % field
capacity between 5 March and 15 May (prior to the water
treatment period) and 8–22 August (recovery period).
Between 30 March and 12 May 2012, all pots were watered
daily with Peters professional ‘Allrounder’ water soluble
NPK fertiliser (20 %, 8.7 %, 16.6 %) plus trace elements
(B, Cu, Fe, Mn, Mo, Zn) at a rate of 1 g l1. Air tem-
peratures were maintained by heating or ventilation at
14
12–16 °C at night and 20–25 °C during the day. The plants
were cut to 50 mm above media level on 4 April, 18 April
and 9 May to ensure all plants were sufficiently established
(82 days after sowing).
Experimental design
Treatments were imposed on 16 May (Day 1; 82 days
after sowing). The experimental design was a 2 9 3 9 3
factorial combination of treatments in a randomised
complete block design with four blocks (19 chicory and
19 plantain pots per block). The three main factors were
plant species (chicory and plantain), water treatment
[optimum (control, 100 % field capacity), dry (63–82 %
field capacity) and very-dry (36–55 % field capacity)]
and defoliation frequency (DF: weekly, fortnightly and
3-weekly) over the period from Day 1 to Day 98. Each
treatment combination was replicated once within each
of the four blocks for each destructive harvest date; Day
1, Day 42 (midpoint of the water treatment period) and
Day 98 (end of the recovery period). This resulted in
eight pots for harvest at Day 1 (two species 9 four
blocks) and 72 pots for each harvest on Day 42 and Day
98 (two species 9 three water treatments 9 three
DF’s 9 four blocks). At defoliation, plants were cut to
50 mm above the potting media level. The length of the
water treatment period (Day 1–84; 12 weeks) was
designed to simulate a severe summer water stress per-
iod. Each individual pot was weighed approximately
every 3 days, and the amount of water required to keep
each pot within its water treatment field capacity range
was calculated and applied. Following the water treat-
ment period, the plants were given a 2-week recovery
period (Day 85–98) during which they received optimum
water (100 % field capacity) and no further defoliation.
Measurements
The volume of water that each pot contained was moni-
tored approximately every 3 days between Day 1 and Day
98, using the time–domain–reflectometry technique (TDR;
CS-615; Campbell Sci., Leicestershire, UK) with moisture
probes to a 20 cm depth. Probes were inserted into the
middle of the pot at each measurement time.
The number of live leaves per plant was recorded at the
time of the pot’s destructive harvest (either at Day 1, 42 or
98). At destructive harvest, taproot diameter at the widest
point and taproot length were recorded and above-ground
herbage mass was dissected into live and dead material.
The root mass was measured after manually washing off
media. All mass data represent the mass per pot (i.e. mass
from four plants) and are presented on a dry weight basis
after oven drying at 60 °C for 48 h.
The shoot mass fraction was calculated as per Hunt
(1978);
© 2015 Blackwell Verlag GmbH, 202 (2016) 13–24

Shoot mass fraction ¼
Total shoot mass per pot
 100
Total shoot mass + total root mass per pot
The relative shoot and root growth rate between each
harvest was calculated as per Hunt (1978);
lnW2  lnW1
Relative Growth Rate (RGR) ¼
time
where: W1 = weight at time 1; W2 = weight at time 2;
Time = number of days between harvest events.
Experiment 2
Plant establishment
The experiment was conducted between 7 January and 16
March 2013. Two plant species, plantain cv. ‘Ceres Tonic’
and chicory cv. ‘Puna II’, were used. The potting medium
consisted of 50 % sand (3 mm) and 50 % aggregate stones
(<10 mm) amended with 90 g short term mix (Woodace,
flowering plant special; LebanonTurf; 14 % N, 6 % P,
11.6 % K + trace elements) and 45 g dolomite for 30 l of
potting medium as a base start fertiliser. The pots were
250 mm in diameter and 250 mm in depth and were filled
with 10.0  0.05 kg of potting medium. Seeds were sown
directly into 40 pots (20 chicory and 20 plantain), on 7 Jan-
uary 2013, and pots were hand watered daily with a sprin-
kler. After seeds germinated, seedlings were thinned to six
per pot and then to four per pot when well established on
12 February 2013 (resulting in a total of 80 chicory plants
and 80 plantain plants).
An automatic watering system incorporating an individ-
ual dripper in each pot was established on 28 January 3013,
and all pots were watered regularly to ensure plant growth
was not limited. All pots were watered daily with liquid
nutrients (1 g l1) (same as in experiment 1) between 27
February and 8 March 2013. Air temperatures were main-
tained by heating or ventilation at 12–16 °C at night and
20–25 °C during the day.
Experimental design
The experiment consisted of a 2 9 2 factorial combination
of treatments in a randomised complete block design with
four blocks. The two main factors were plant species (chic-
ory and plantain) and water treatment [optimum (control)
and very-dry]. The optimum water treatment provided a
set volume of water per pot (same for chicory and plantain)
which was based on the level of sunlight and plant growth,
to ensure soil water content was not limiting plant growth.
The very-dry water treatment consisted of the plants receiv-
ing no water. The water treatment period started on 11
March (Day 1) and lasted for 5 days (days 1–5). Each treat-
ment combination was replicated once within each of the
© 2015 Blackwell Verlag GmbH, 202 (2016) 13–24
Responses of Plantago and Cichorium to Water Stress
four blocks for each measurement group (destructive mea-
surements and non-destructive measurements).
Measurements
All physiological plant measurements were carried out
during days 1–5 of the treatment period during peak
sunlight hours (between 12 and 2 pm daily). Measure-
: ments were collected in a block by block manner. The
photosynthetic characteristics of the plants [photosyn-
thetic rate (Pn), evapotranspiration rate (E) and sto-
matal conductance (Cs)] were studied using a LI–6400
portable photosynthetic system (made by LI–COR Bio-
sciences, Lincoln, Nebraska, USA). One measurement
was taken per pot (on one leaf of one plant) on the
most recent fully expanded leaf of the set of non-
destructive plants. Water potential (Ψw) was measured
using a Scholander pressure chamber as described by
Turner (1988). Ψw measurements were undertaken on
the set of destructive measurement pots (one measure-
ment per pot from one plant). Destructive harvest
measurements on Day 1 and Day 6 were undertaken
as per Experiment 1. The volume of water that each
pot contained was monitored daily using the TDR
technique as in Experiment 1.
Experiment 3
Site
The experiment was conducted on 11 February 2013 at
the Massey University Pasture and Crop Research Unit,
5 km south of Palmerston North, New Zealand. January
and February are typically the driest months of the year
at this site, and 2013 represented a particularly dry sum-
mer (29 and 34 mm of rain during January and February
of 2013 compared to 10-year average rainfall rates of 52
and 55 mm, respectively). The soil type was a Manawatu
fine sandy loam (Dystric Fluventic Eutrochrept, Hewitt
1988), with pH 5.7. An existing paddock with a sward
mix based on chicory cv. ‘Puna II’ and plantain cv.
‘Ceres Tonic’ was used (Cave et al. 2015). The paddock
had been sown in March 2011, and thus, the plants were
in their second growing season at the time of the experi-
ment. The paddock had been regularly grazed with sheep,
from a pre-grazing height of 25 cm to a post-grazing
height of 5 cm.
Experimental design
Twenty six pairs of plants (one chicory and one plantain,
n = 52 in total) were excavated so that all root material
was intact. Plants within each pair were within a 50 cm
radius of each other. Roots were washed so that all visible
dirt was removed and cut into 10 cm rooting intervals
(0–10 and 10–20 cm). No roots were deeper in the soil
15
Cranston et al.
than 20 cm. Roots were then oven dried at 60 °C for 48 h,
and dry weight was recorded.
Statistical analysis
All data were analysed using the SAS mixed data procedure
(SAS Version 9.2; SAS Institute Inc., Cary, NC, USA). Only
significant interactions (P ≤ 0.05) were presented in the
results section. When significant (P ≤ 0.05) effects of treat-
ments were detected, mean separation was conducted by
the PDIFF option in SAS.
In Experiment 1, data were analysed separately each
destructive harvest period (Day 1, Day 42 and Day 98) and
for each parameter (absolute shoot mass, relative shoot
growth rate, taproot diameter, root length, absolute root
mass, relative root growth rate, shoot mass fraction, num-
ber of live leaves per plant and leaf mass). At Day 1 (Har-
vest 1), the model included the fixed effects of plant species
(chicory and plantain) and the random effect of block. At
Day 42 (Harvest 2) and Day 98 (Harvest 3), each model
included the fixed effects of plant species (chicory and
plantain), water treatment (optimum and dry and very-
dry) and DF (weekly, fortnightly and 3-weekly) and all
two-way and three-way interactions between plant species,
water treatment and DF and the random effect of block.
In Experiment 2, data were analysed separately each
destructive harvest period (Day 1 and Day 6) and for each
parameter. At Day 1 (Harvest 1), the model included the
fixed effects of plant species (chicory and plantain) and the
random effect of block. At Day 6 (Harvest 2), the model
included the fixed effects of plant species (chicory and
plantain), water treatment (optimum and very-dry) and
the interaction between plant species and water treatment
and the random effect of block.
The physiological plant measurement data (Pn, E, Cs
and Ψw) and soil volumetric water content were also
analysed using the MIXED procedure in SAS. Each
model included the fixed effects of plant species (chicory
and plantain) and water treatment (optimum and very-
dry) and the interaction between plant species and water
treatment and included day within block as a repeated
measure.
In Experiment 3, t-tests were used to compare the mass
of root within the depth interval within each species. The
data representing the mass of roots within each depth
interval between species were not normally distributed and
were analysed using a nonparametric ANOVA (Kruskal–Wal-
lis Test).
Results
In all sections, interactions are only presented and stated
where they were found to be significant (P ≤ 0.05).
16
Experiment 1
Water treatment
All water treatments were successfully established by the 31
May, such that the average soil volumetric contents were
26.7  0.2, 26.3  0.2, 8.6  0.2, 8.2  0.2, 6.4  0.2,
6.2  0.2 m3 m3 for chicory-optimum, plantain-opti-
mum, chicory-dry, plantain-dry, chicory-very-dry and
plantain-very-dry, respectively.
Day 1 – Harvest 1
Chicory had a greater (P ≤ 0.05) taproot diameter (11.9 vs.
4.7  0.4 mm) and absolute root mass (4.2 vs. 1.5 
0.5 g) than plantain. Plantain had a greater (P ≤ 0.05)
root length (174 vs. 148  17 mm), number of live leaves
per plant (44 vs. 25  3) and shoot mass fraction (83 vs.
61  2 %) than chicory.
Day 42 – Harvest 2
Above-ground herbage measures. Plantain had a greater
(P ≤ 0.05) absolute shoot production, relative shoot
growth rate (days 1–42), shoot mass fraction and number
of live leaves per plant than chicory (Table 1). Chicory had
a greater (P ≤ 0.05) leaf mass than plantain. Absolute shoot
production and relative shoot growth rate were greater
(P ≤ 0.05) under the 3-weekly DF than the fortnightly DF,
which was in turn greater (P ≤ 0.05) than the weekly DF
(Table 1).
Below-ground herbage measures. There were significant
two-way interactions (P ≤ 0.05) between species and water
treatment and between species and DF for taproot diameter
(Table 2). Plantain taproot diameter did not differ
(P > 0.05) across water treatments but was lower
(P ≤ 0.05) than all chicory by water treatment combina-
tions. Chicory taproot diameter was greater (P ≤ 0.05)
under optimum water treatment than under dry and very-
dry water treatments, which did not differ (P > 0.05).
Plantain taproot diameter did not differ (P > 0.05) across
DF’s. Chicory taproot diameter was greater (P ≤ 0.05)
under three-weekly DF than under weekly and fortnightly
DF’s, which did not differ (P > 0.05).
There was a two-way interaction (P ≤ 0.05) between
water treatment and DF for absolute root mass (Table 2).
Under three-weekly DF, optimum plants had a greater
(P ≤ 0.05) absolute root mass than dry and very-dry plants,
which did not differ (P > 0.05). Whereas under both
weekly and fortnightly DF’s, water treatment had no effect
(P > 0.05) on absolute root mass. Chicory had a greater
(P ≤ 0.05) absolute root mass than plantain (4.9 vs.
3.8  0.4 g).
Plantain had a greater (P ≤ 0.05) relative root growth
rate (days 1–42) than chicory (0.02 vs. 0.00  0.002
g g1 day1). Three-weekly DF plants had a greater
© 2015 Blackwell Verlag GmbH, 202 (2016) 13–24
 Responses of Plantago and Cichorium to Water Stress

Table 1 Effect of plant species (plantain and chicory), water treatment (optimum, dry and very-dry) and defoliation frequency (DF: weekly, fort-
nightly and 3-weekly) at Day 42 on absolute shoot mass, relative shoot growth rate (days 1–42), shoot mass fraction (Shoot/(shoot + root) 9 100),
number of live leaves per plant and leaf mass in Experiment 1

Absolute shoot Relative shoot Shoot mass Number of live Leaf mass
mass (g) growth rate (g g1 day1) fraction (%) leaves per plant (g leaf1)

Species
Plantain 14.7 b 0.016 b 57.4 b 35.2 b 0.032 a
Chicory 11.8 a 0.013 a 40.3 a 20.1 a 0.037 b
S.E.M. 0.6 0.001 1.8 1.3 0.002
Water
Optimum 13.9 0.015 45.6 a 30.1 b 0.031 a
Dry 13.6 0.015 52.4 b 28.8 b 0.036 b
Very-dry 12.3 0.013 48.6 ab 24.0 a 0.037 b
S.E.M. 0.7 0.001 2.1 1.4 0.002
DF
Weekly 10.8 a 0.010 a 53.7 b 28.2 0.026 a
Fortnightly 13.2 b 0.014 b 51.6 b 27 0.041 c
Three-weekly 15.7 c 0.019 c 41.4 a 27.8 0.037 b
S.E.M. 0.7 0.001 2.1 1.4 0.002

Data presented as means  S.E.M. Means within columns and main effects having different letters are significantly different at P ≤ 0.05.

(P ≤ 0.05) relative root growth rate (days 1–42) than fort-
nightly DF plants, which in turn had a greater (P ≤ 0.05)
relative root growth rate than weekly DF plants (0.019 vs.
0.010 vs. 0.008  0.002 g g1 day1).
Day 98 – Harvest 3
Above-ground herbage measures. All plants survived the
entire experimental period. There was a three-way interac-
tion (P ≤ 0.05) between species, water treatment and DF
for relative shoot growth rate (days 42–98). Plantain had a
greater (P ≤ 0.05) relative shoot growth rate than chicory
under all water treatment by DF combinations (Fig. 1a).
Under all water treatments, weekly DF resulted in a lower
(P ≤ 0.05) relative shoot growth rate than less frequent
DF’s.
There was a two-way interaction (P ≤ 0.05) between water
treatment and DF on shoot mass grown during the recovery
period. Very-dry plants under fortnightly and 3-weekly DF’s
had a greater (P ≤ 0.05) absolute shoot mass grown during
the recovery period than dry and optimum plants, which did
not differ (P > 0.05) (Fig. 2). Under weekly DF’s dry plants
had a greater (P ≤ 0.05) absolute shoot mass grown during
the recovery period than both optimum and very-dry plants,
which did not differ (P > 0.05). Plantain had a greater
(P ≤ 0.05) absolute shoot mass grown during the recovery
period than chicory (Table 3).
Absolute shoot mass grown during the water treatment
period did not differ (P > 0.05) between chicory and
plantain (Table 3). Very-dry plants had a lower (P ≤ 0.05)
absolute shoot mass grown during the water treatment per-
iod than optimum and dry plants, which did not differ
(P > 0.05). Three-weekly DF plants had a greater (P ≤
0.05) absolute shoot mass grown during the water treat-
ment period than weekly and fortnightly DF plants, which
did not differ (P > 0.05). Plantain had a greater (P ≤ 0.05)
shoot mass fraction and number of live leaves per plant
and a lower (P ≤ 0.05) leaf mass than chicory (Table 3).
Below-ground herbage measures. There was a two-way
interaction (P ≤ 0.05) between species and DF for absolute
root mass. Within chicory, the 3-weekly DF had a greater
(P ≤ 0.05) absolute root mass than the weekly and fort-
nightly DF’s, which did not differ (P > 0.05) (3.8 vs. 4.7 vs.
7.5  0.4 for weekly, fortnightly and 3-weekly DF’s,
respectively). Similarly, within plantain, the 3-weekly DF
had a greater (P ≤ 0.05) absolute root mass than the weekly
DF, with fortnightly DF not differing (P > 0.05) from
either treatment (1.6 vs. 2.2 vs. 2.8  0.4 for weekly, fort-
nightly and 3-weekly DF’s, respectively). Across all DF’s
chicory had a greater (P ≤ 0.05) absolute root mass than
plantain.
There was a three-way interaction (P ≤ 0.05) between
species, water treatment and DF for relative root growth
rate (days 42–98) (Fig. 1b). In the very-dry water treat-
ment, chicory always had a greater (P ≤ 0.05) relative root
growth rate than plantain. However, this pattern was not
always observed under the optimum and dry water treat-
ment whereby, chicory had a lower (P ≤ 0.05) relative root
growth rate under fortnightly DF than plantain.
Chicory had a greater (P ≤ 0.05) taproot diameter than
plantain (11.6 vs. 5.0  0.2 mm). Water treatment had
no effect (P > 0.05) on taproot diameter (data not
shown). Taproot diameter was greater (P ≤ 0.05) under
3-weekly DF than under weekly DF, while fortnightly DF

© 2015 Blackwell Verlag GmbH, 202 (2016) 13–24 17

Cranston et al.

Table 2 Effect of plant species (plantain and chicory), water treatment Physiological measurements
(optimum, dry and very-dry) and defoliation frequency (DF: weekly, There were three-way interactions (P ≤ 0.05) between
fortnightly and 3-weekly) at Day 42 on taproot diameter, root length
plant species, water treatment and day of measurement for
and absolute root mass in Experiment 1
photosynthetic rate, evapotranspiration rate and leaf water
Taproot Root Absolute root potential. On Day 1, the photosynthetic rate (Pn) within
diameter (mm) length (mm) mass (g) each plant species did not differ (P > 0.05) under optimum
Species 9 water
and very-dry water treatments; however, plantain had a
Plantain-optimum 4.1 a greater (P ≤ 0.05) Pn than chicory (Fig. 4a). Between days
Plantain-dry 4.2 a 2 and 5, the Pn of chicory and plantain under optimum
Plantain-very-dry 4.1 a water treatment did not differ (P > 0.05) and was greater
Chicory-optimum 12.7 c (P ≤ 0.05) than the Pn of plants under very-dry water
Chicory-dry 11.6 b treatment. On days 2 and 4, plantain-very-dry had a greater
Chicory-very-dry 11.1 b
(P ≤ 0.05) Pn than chicory-very-dry. While on days 3 and
S.E.M. 0.3
Species 9 DF
5, the Pn of plantain-very-dry did not differ (P > 0.05) to
Plantain-weekly 4.2 a 239 b chicory-very-dry.
Plantain-fortnightly 4.1 a 257 c On Day 1, the evapotranspiration rate (E) within each
Plantain-3-weekly 4.1 a 284 d plant species did not differ (P > 0.05) under optimum and
Chicory-weekly 11.4 b 135 a very-dry water treatments; however, plantain had a greater
Chicory-fortnightly 11.5 b 140 a (P ≤ 0.05) E than chicory (Fig. 4b). Between days 2 and 5,
Chicory-3-weekly 12.4 c 136 a
the E did not differ (P > 0.05) between species but was
S.E.M. 0.3 8
Water 9 DF
greater (P ≤ 0.05) for optimum water treatment plants
Optimum-weekly 183 a 2.5 a than very-dry water treatment plants.
Dry-weekly 181 a 2.2 a On Day 1, there were no differences (P > 0.05) in the
Very-dry-weekly 197 abc 2.8 ab leaf water potential (Ψw) between species or water treat-
Optimum-fortnightly 188 a 4.6 bc ments (Fig. 4c). Between days 2 and 5, the Ψw of chicory-
Dry-fortnightly 217 d 4.7 c optimum and plantain-optimum did not differ (P > 0.05)
Very-dry-fortnightly 191 ab 4 abc
and were lower (P ≤ 0.05) than the Ψw of very-dry water
Optimum-3-weekly 213 cd 8.2 d
Dry-3-weekly 209 bcd 4.7 c
treatment plants. The Ψw of chicory-very-dry was greater
Very-dry-3-weekly 208 bcd 5.2 c (P ≤ 0.05) than plantain-very-dry on Day 3 and did not
S.E.M. 9 0.7 differ (P > 0.05) on days 2, 4 and 5.
Plantain tended (P = 0.09) to have greater stomatal con-
Data presented as means  S.E.M.; only significant results are shown.
ductance (Gs) than chicory (2760 vs. 982  499 m
Means within columns and treatments having different letters are sig-
nificantly different at P ≤ 0.05.
mol m2 s1). Very-dry water treatment plants tended
(P = 0.06) to have lower Gs than optimum water treat-
ment plants (758 vs. 2985  499 m mol m2 s1).
did not differ (P > 0.05) from either (8.0 vs. 8.3 vs.
8.6  0.2 mm under weekly, fortnightly and 3-weekly
Experiment 3
DF’s, respectively).
Both chicory and plantain had a greater (P ≤ 0.05) root
mass in the 0–10 cm soil depth profile than in the
Experiment 2
10–20 cm soil depth profile (Table 4). The root mass in
Water treatment the 0–10 cm soil depth profile did not differ (P > 0.05)
There was a three-way interaction (P ≤ 0.05) between plant between chicory and plantain. Chicory had an 18 % greater
species, water treatment and day of measurement (Fig. 3). (P ≤ 0.05) root mass in the 10–20 cm soil depth profile
From Day 3 onwards, the soil volumetric water content of than plantain.
the very-dry water treatment was lower (P ≤ 0.05) than the
optimum water treatment and did not differ (P > 0.05)
Discussion
between species. Within the optimum water treatment, the
soil volumetric water content did not differ (P > 0.05)
Morphological differences between chicory and plantain
between species from Day 4 onwards. Consequently, by
Day 6, the optimum water treatment had a greater The root of chicory consists of a solid deep taproot (Kemp
(P ≤ 0.05) absolute shoot mass than the very-dry water et al. 2002). Conversely, while plantain is technically tap-
treatment (39.4 vs. 29.9  1.3 g). rooted, its root system is more fibrous due to a greater

18 © 2015 Blackwell Verlag GmbH, 202 (2016) 13–24


(a)
(b)
Fig. 1 Effect of species [chicory (black) and
plantain (grey bars)], water treatment [opti-
mum (Opt), dry and very-dry (VD)] and defolia-
tion frequency [weekly (1), fortnightly (2) and
3-weekly (3)] on the relative shoot growth rate
(a; S.E.M. = 0.002) and the relative root
growth rate (b; S.E.M. = 0.003) between days
42 and 98 in Experiment 1.
number of adventitious roots compared to chicory (San-
derson et al. 2003). Therefore, as expected in experiment 1,
chicory consistently had a greater root mass and taproot
diameter than plantain. Furthermore, at the end of experi-
ment 1, under very-dry conditions, the difference in the
relative root growth rate of chicory compared to plantain
was generally greater than that under less severe water
treatments. These root differences suggest that chicory may
have an advantage over plantain under very-dry conditions
and may be more persistent in environments which experi-
ence severe droughts. Plant survival in drought periods has
been shown to be related to root density and depth (Volaire
and Thomas 1995, Bahrani et al. 2010, Wedderburn et al.
© 2015 Blackwell Verlag GmbH, 202 (2016) 13–24
Responses of Plantago and Cichorium to Water Stress
2010). Furthermore, a deep root system with a high density
of roots at depth is a major trait to sustain a higher shoot
yield in water-limited environments (Carrow 1996, Wasson
et al. 2012, White and Snow 2012). Under glasshouse con-
ditions, the roots of chicory were limited by pot depth and
likely limiting its potential advantage for water extraction
compared to plantain. In experiment three, under field
conditions, chicory had a greater mass of roots in the dee-
per 10–20 cm soil horizon than plantain. Under field con-
ditions, where root length is not limited Brown et al.
(2005) found 1-year-old chicory was able to extract water
to a maximum depth of 1.9 m. While, Nie et al. (2008)
found 3-year-old plantain had a mean rooting depth of
19
Cranston et al.

as an adaptive feature for plant survival, because it allows

Fig. 2 Effect of water treatment [optimum (black), dry (grey) and very-
dry (white bars)] and defoliation frequency [weekly (1), fortnightly (2)
and 3-weekly (3)] on absolute shoot mass grown during the recovery
period (days 85–98), in Experiment 1. Vertical lines above the bars rep-
resent S.E.M.
0.97 m, with a substantially greater root density in the top-
soil (0–10 cm) than in the subsoil (10–110 cm).
In experiment one, plantain consistently had a greater
relative shoot growth rate than chicory. In a preliminary
experiment, Cranston (2014) also showed plantain had a
greater relative shoot growth rate than chicory. Van Andel
and Biere (1990) suggested that relative shoot growth rate
is a useful indicator of plant productivity. Previous research
under field conditions has shown that plantain has a
greater herbage yield than chicory under optimum growth
conditions (Powell et al. 2007). However, when subjected
to water stress, slower herbage growth has been suggested
plants to divert assimilates and energy, otherwise used for
shoot growth, to maintain root growth, improving water
acquisition (Chaves et al. 2003). Therefore, it could be
expected that the slower relative shoot growth rate of chic-
ory compared to plantain would be more pronounced
under the very-dry water treatment, but this was not found
to be the case. This may therefore suggest that chicory sim-
ply has a slower shoot growth rate than plantain.
In experiment one, plantain consistently had a greater
shoot mass fraction (shoot/shoot + root) than chicory, as
found by Cranston (2014). It is well established that chic-
ory allocates a greater energy reserve to root mass com-
pared to plantain which focusses on shoot production
(Alemseged 2000, Sanderson and Elwinger 2000, Powell
et al. 2007). Alemseged (2000) reported the shoot to root
ratio in chicory steadily declined over time with growth as
it allocated more energy to root than shoot, probably in
response to localised nutrient depletion. Consequently,
Alemseged (2000) concluded that this high degree of plas-
ticity may enable chicory to sustain a high rate of resource
capture by placing its roots in the resource-rich zone of the
soil profile. Presumably, chicory’s lower shoot mass frac-
tion would help it to survive and persist under severe water
stress. Conversely, in plantain, it was postulated that under
severe water stress, the greater partitioning of energy to
above-ground growth would not be maintained, or alterna-
tively that plant survival may be negatively affected. How-
ever, this was not the case as both species fully recovered
following the 12-week period of water stress, in experiment
one. This illustrates that both plantain and chicory display
a degree of resilience in terms of recovering from soil water

Table 3 Effect of plant species (plantain and chicory), water treatment [optimum, dry and very-dry (VD)] and defoliation frequency (DF; weekly, fort-
nightly and 3-weekly) on shoot mass grown during water treatment period and during recovery period, shoot mass fraction (Shoot/
(shoot + root) 9 100), number of live leaves per plant and leaf mass at Day 98 in Experiment 1

Absolute shoot mass grown Absolute shoot mass Shoot mass Number of live Leaf mass
during water treatment period (g) grown during recovery period (g) fraction (%) leaves per plant (g leaf1)

Species
Plantain 14.8 4.2 b 66.2 b 33.8 b 0.032 a
Chicory 13.7 3.5 a 40.2 a 24.5 a 0.036 b
S.E.M. 0.6 0.3 1.8 0.9 0.002
Water
Optimum 15.7 b 3.2 a 50.3 32.0 b 0.025 a
Dry 14.8 b 3.8 b 54.7 27.7 a 0.035 b
VD 12.2 a 4.6 c 54.5 27.8 a 0.042 c
S.E.M. 0.7 0.3 2.1 1.0 0.002
DF
Weekly 12.5 a 3.8 60.4 b 30.2 0.032 a
Fortnightly 13.8 a 3.7 51.9 a 29.0 0.032 a
Three-weekly 16.3 b 4.1 47.3 a 28.3 0.038 b
S.E.M. 0.7 0.3 2.1 1.0 0.002

Data presented as means  S.E.M. Means within columns and main effects having different letters are significantly different at P ≤ 0.05.

20 © 2015 Blackwell Verlag GmbH, 202 (2016) 13–24

 Responses of Plantago and Cichorium to Water Stress

(a)

(b)
Fig. 3 Change in soil volumetric water content (m3 m3) of the four
species by water treatment combinations: chicory-optimum (●), plan-
tain-optimum (○), chicory-very-dry (▼) and plantain-very-dry (M) over
the 5-day treatment period, in Experiment 2.

stress. Previous research has also shown chicory and plan-

tain can survive periods of water stress (Mook et al. 1989,
Brown et al. 2003, Nie et al. 2008).

Effect of defoliation frequency and water treatment

Carbohydrate reserves in perennial forage plants are an
important source of energy necessary for winter survival (c)
and initiation of growth in spring (Kim et al. 1991, Hen-
dershot and Volenec 1993). Furthermore, they can provide
the necessary energy to support shoot growth following
defoliation (Kim et al. 1993). Li et al. (1997a) compared
three chicory cultivars and found that taproot diameter
reflected total root reserves, suggesting that taproot diameter
can be used as an indicator of root carbohydrate reserves.
At Day 42 of experiment one, defoliation frequency had no
effect on the taproot diameter of plantain; however, the tap-
root diameter of chicory was greater under 3-weekly defoli-
ation than under more frequent defoliation. By Day 98 of
experiment one, defoliation frequency had the same effect
on both chicory and plantain with plants under 3-weekly
defoliation having a greater taproot diameter than those Fig. 4 Effect of species [chicory (black), plantain (white)] and water
under weekly defoliation. These results suggest that chicory treatment [optimum (circle) and very-dry (triangle)] on the photosyn-
and plantain would be more likely to persist in the long thetic rate (a), evapotranspiration rate (b), leaf water potential (c) of
term under 3-weekly defoliation than weekly defoliation, plants during the water treatment period (days 1–5), in Experiment 2.
given their greater root reserves. Furthermore, the taproot
diameter of chicory showed a response to defoliation fre- Experiment one found the relative shoot and root
quency earlier than plantain, suggesting plantain, in the growth rates of both chicory and plantain were reduced
short term at least, might be more resilient to defoliation under more frequent defoliation. This has previously been
frequency than chicory. Somewhat in support of this, Yu reported in chicory under field conditions (Li et al. 1997b,
et al. (2008) reported the taproot diameter of plantain was Belesky et al. 1999, Alemseged et al. 2003, Sanderson et al.
unaffected by defoliation height, whereas that of chicory 2003). In plantain, a 5-week defoliation interval has been
was reduced under severe defoliation (lower residual), indi- shown to produce greater shoot dry matter than a 3-week
cating plantain is less sensitive to defoliation than chicory. defoliation interval (Sanderson et al. 2003, Labreveux et al.

© 2015 Blackwell Verlag GmbH, 202 (2016) 13–24 21

Cranston et al.
Table 4 Experiment 3: Effect of species (chicory and plantain) on the
average root mass per plant (g DM) at 0–10 cm and 10–20 cm, in
Experiment 3
Root mass per plant (g DM)
Soil depth Plantain Chicory
0–10 cm 0.91 b  0.14 1.20 b  0.17
10–20 cm 0.14 a†  0.06 0.20 a¥  0.03
Data presented as means  S.E. Means within columns having differ-
ent letters are significantly different at P ≤ 0.05. Means within rows
having different symbols (†, ¥) are significantly different at P ≤ 0.05.
2004). While in Treinta y Tres, Uruguay, an area typically
hotter than Palmerston North, New Zealand, Ayala et al.
(2011) found a 3-week defoliation interval for plantain
produced more dry matter than a 6-week defoliation inter-
val. Yu et al. (2008) found the shoot dry matter of chicory
and plantain increased with decreasing mowing frequency,
between one and three weeks. In experiment 1, 3-weekly
defoliation consistently resulted in a greater shoot mass
and root mass compared to more frequent defoliation.
Combined these results suggest for both chicory and plan-
tain, 3-weekly defoliation is likely to produce a greater
herbage yield and a more persistent sward than a more fre-
quent defoliation regimen.
In experiment 1, both chicory and plantain, under fort-
nightly and 3-weekly defoliation, subjected to the very-dry
water treatment produced a greater shoot mass during the
recovery period (compensatory growth) than the dry and
optimum water treatments. While under weekly defolia-
tion, dry water treatment plants grew greater shoot mass
during the recovery period than the very-dry and optimum
water treatments plants. It is likely that under severe
drought and weekly defoliation, root reserves are dimin-
ished and when the water stress is removed the plant does
not have sufficient root reserves to display compensatory
growth. While under a mild drought (i.e. the dry treatment)
and weekly defoliation, the plant maintains sufficient root
reserves to display compensatory growth. Similarly, under
less frequent defoliation, the plant should be able to display
compensatory growth following a severe drought. Thus,
these results suggest that both plantain and chicory have the
potential to produce dry matter, in the short term at least,
following a period of severe drought, provided that the
defoliation interval has been a minimum of 2 weeks.
Physiological differences between chicory and plantain
In experiment 2, plantain and chicory generally displayed
similar rates of photosynthesis and evapotranspiration per
unit of leaf area under optimum and very-dry conditions.
Photosynthesis and evapotranspiration rates of both species
22
were lower under very-dry conditions than optimum con-
ditions. However, experiment one showed plantain consis-
tently had a greater number of live leaves per plant and a
lighter mass per leaf than chicory. This difference suggests
that chicory and plantain have different growth strategies
with plantain investing its energy in many smaller leaves
while chicory supports fewer larger leaves. Combined, these
results suggest plantain’s ability to yield more than chicory
under water stress conditions in the glasshouse could be as
a result of morphological factors which enable greater light
interception (a growth strategy suggested by Korner
(1991)) rather than physiological differences between the
two species. However, further research is required to con-
firm this.
Conclusions
In conclusion, under simulated drought conditions plan-
tain generally had superior shoot growth, whereas chicory
displayed superior root growth. Neither chicory nor plan-
tain responded well to weekly defoliation and performed
best under 3-weekly defoliation. Plantain and chicory dis-
played similar rates of photosynthesis and evapotranspira-
tion per unit of leaf area under water stress conditions. In
the field, chicory had roots deeper in the soil profile than
plantain; further supporting evidence to suggest chicory is
better than plantain for very-dry environments.
Acknowledgements
The excellent technical assistance of Kay Sinclair and Bob-
bie Cave as well as PGU staff; Steve Ray, Lindsay Sylva and
Lesley Taylor are gratefully acknowledged. The senior
author is thankful for the financial assistance of MacMillan
Brown Agricultural Research Scholarship, John Hodgson
Pastoral Science Scholarship, Helen E Akers PhD Scholar-
ship, New Zealand Federation of Graduate Women Post-
Graduate Fellowship and a Massey Doctoral Scholarship.
P.R.Kenyon was partially funded by the National Centre
for Growth and Development (Gravida).
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