Crop Ecology & Physiology

Daytime Sprinkler Irrigation Effects on Net Photosynthesis

of Maize and Alfalfa
Yenny F. Urrego-Pereira, Antonio Martínez-Cob, Victoria Fernández, and José Cavero*

ABSTRACT
During sprinkler irrigation some water is lost due to drift and evaporation. After irrigation, plant-intercepted water is lost due to
evaporation. The water loss causes microclimatic changes, which may involve positive or negative plant physiological responses.
We studied the changes in net photosynthesis of maize (Zea mays L.) and alfalfa (Medicago sativa L.) associated with irrigation
with a solid-set sprinkler system. For each species, measurements were made simultaneously in two plots, one being irrigated and
the other not being irrigated. Two automatic canopy chambers connected to two CO2 infrared gas analyzers were used. Sprinkler
irrigation decreased air temperature (1.5°C on maize, 1.7°C on alfalfa), air vapor pressure deficit (VPD) (0.44 kPa for both crops)
and canopy temperature (5.1°C on maize, 5.9°C on alfalfa). Sprinkler irrigation decreased maize net photosynthesis on 80% of
the days and the mean reduction was 19%. Sprinkler irrigation increased alfalfa net photosynthesis on 36% of days, decreased it
on 14% of days, and had no effect on half of the days. The decrease of maize net photosynthesis during sprinkler irrigation was
linked to the high leaf wettability (water contact angles from 60–80°) and the decrease in temperature below the optimum range
for photosynthesis. The higher hydrophobicity of alfalfa leaves (water contact angles >120°) and the wide range of optimum
temperature for alfalfa photosynthesis may be the reasons why photosynthesis remained unaffected by sprinkler irrigation. The
results suggest that daytime sprinkler irrigation with solid-set should be avoided for maize but can be used for alfalfa.

D uring a sprinkler irrigation event, some water is

lost due to wind drift and evaporation that occurs as
water travels from the sprinkler nozzles to the crop, and to
evaporation that occurs for water intercepted by stems and
leaves after the irrigation event (Tolk et al., 1995). The water
that is evaporated produces microclimatic changes: decreases
the temperature and VPD of the air (Robinson, 1970; Thomp-
son et al., 1993; Tolk et al., 1995; Cavero et al., 2009). These
changes are greater during daytime (Cavero et al., 2009).
The microclimatic changes during daytime sprinkler
irrigation cause physiological changes in the crops, which
could affect their growth and yield. A relevant physiological
change during sprinkler irrigation is the reduction in crop
transpiration (McNaughton, 1981; Tolk et al., 1995; Martínez-
Cob et al., 2008), which is considered positive because it
represents a reduction of wind drift and evaporation losses
(WDEL) (Martínez-Cob et al., 2008). Another major
physiological change during sprinkler irrigation is the decrease
in crop canopy temperature (Steiner et al., 1983; Tolk et al.,
1995; Saadia et al., 1996; Cavero et al., 2009), which could
have a positive effect on photosynthesis when leaf temperatures
are too high (Mahan et al., 1995; Wanjura and Upchurch,
Y.F. Urrego-Pereira, A. Martínez-Cob, J. Cavero, Dep. Soil and Water, Estación
Experimental de Aula Dei (CSIC), Avda. Montañana 1005, 50059 Zaragoza,
Spain; V. Fernández, Forest Genetics and Ecophysiology Research Group,
School of Forest Engineering, Technical University of Madrid, 28040 Madrid,
Spain. Received 8 Mar. 2013. *Corresponding author (jcavero@eead.csic.es).
Published in Agron. J. 105:1515–1528 (2013)
doi:10.2134/agronj2013.0119
Copyright © 2013 by the American Society of Agronomy, 5585 Guilford
Road, Madison, WI 53711. All rights reserved. No part of this periodical may
be reproduced or transmitted in any form or by any means, electronic or
mechanical, including photocopying, recording, or any information storage
and retrieval system, without permission in writing from the publisher.
2000); for instance, during the afternoon of summer months in
semiarid climates. However, if leaf temperature decreases below
an optimum value, photosynthesis could also decrease (Mahan
et al., 1995).
Hirasawa and Hsiao (1999) found a decrease in stomatal
conductance and photosynthesis of maize during the
afternoon in a well irrigated maize crop in California (high
evapotranspiration). This decline in photosynthesis during
the afternoon of summertime days has been observed in other
studies (Huck et al., 1983; Puech-Suanzes et al., 1989; Bunce,
1990a, 1990b; Pettigrew et al., 1990). Hirasawa and Hsiao
(1999) indicated that the decrease in photosynthesis occurred
because the plants could not transpire at the rate imposed by
the atmospheric conditions, which caused a decrease in leaf
water potential (LWP). Hsiao (1990) proposed that sprinkler
irrigation could reduce the water stress experienced by the
plant due to microclimatic changes during sprinkling. Howell
et al. (1971) found that intermittent mist irrigation increased
the LWP of southern pea [Vigna unguiculata (L.) Walp.]
during irrigation and led to a higher yield than the non-mist-
irrigated treatment. Cavero et al. (2009) found that sprinkler
irrigation increased the LWP of maize from –1.2 and –1.4 MPa
to –0.54 MPa. Given that photosynthesis of crops increases
as LWP increases (Boyer, 1970a, 1970b; Beadle et al., 1973),
sprinkler irrigation could result in increased photosynthesis.
However, Cavero et al. (2008) reported a 10% reduction in
maize grain yield with daytime solid-set sprinkler irrigation
compared with nighttime irrigation.
Abbreviations: CU, Christiansen coefficient of uniformity; ETo, reference
evapotranspiration; ETc, crop evapotranspiration; FC, field capacity; GMT,
Greenwich Mean Time; IRGA, infrared gas analyzer; LWP, leaf water
potential; PAR, photosynthetically active radiation; VPD, vapor pressure
deficit; WDEL, wind drift and evaporation losses; WP, wilting point.

A g ro n o my J o u r n a l  •  Vo l u m e 10 5 , I s s u e 6  •  2 013 1515
Table 1. Soil characteristics of the experimental fields.
Depth pH C N CaCO3 Sand Silt Clay FC† WP‡
m —————————————————– % —————————————————– ———– m3 m–3————
Exp. 1
0.0–0.3 8.4 0.86 0.110 30.9 26.5 45.4 28.1 0.351 0.197
0.3–0.6 8.4 0.72 0.102 31.6 24.0 46.9 29.1 0.351 0.217
0.6–0.9 8.4 0.44 0.088 30.7 17.4 50.0 32.6 0.344 0.196
0.9–1.2 8.6 0.38 0.075 30.3 19.1 50.3 30.6 0.329 0.171
Exp. 2
0.0–0.3 8.1 0.82 0.070 36.0 51.0 35.5 13.4 0.269 0.096
0.3–0.6 8.2 0.52 0.045 39.4 54.4 33.8 11.8 0.250 0.083
0.6–0.9 8.3 0.38 0.036 38.2 56.4 32.8 10.8 0.243 0.071
0.9–1.2 8.4 0.30 0.028 38.8 55.8 34.4 9.7 0.243 0.065
† FC, field capacity (–0.033 MPa).
‡ WP, wilting point (–1.5 MPa).

MATERIALS AND METHODS

Several studies have found that water from rainfall, fog, dew,
or artificial misting can affect the photosynthesis of plants
(Smith and McClean, 1989; Brewer and Smith, 1994, 1995;
Ishibashi and Terashima, 1995; Hanba et al., 2004). It has
been shown that the effect of leaf wetting on photosynthesis
depends on leaf wettability. A material is defined as wettable
when contact angles of drops deposited on to its surface are
below 90° and non-wettable when angles are above 90°. The
solid–liquid interactions between drops of water deposited on
to plant surfaces will depend both on the physical structure
(i.e., micro- and nanoroughness) and chemical composition
of every particular surface (Brewer et al., 1991; Khayet and
Fernández, 2012). Photosynthesis increased or was not
affected by wetting when leaves were non-wettable (Smith and
McClean, 1989; Hanba et al., 2004), but seemed to decrease
when leaves were wettable (Smith and McClean, 1989; Brewer
and Smith, 1994, 1995; Ishibashi and Terashima, 1995; Hanba
et al., 2004). This different behavior is related to the negative
effect on photosynthesis (lower CO2 uptake) of the water
deposited on to the surface of wettable leaves that may limit gas
exchange, in contrast to the positive effect for photosynthesis
of the lower VPD of the air when leaves are non-wettable
(Smith and McClean, 1989; Brewer et al., 1991; Brewer and
Smith, 1995, 1997). The relationship between leaf wettability
and photosynthesis has been studied mainly in natural
ecosystems and in relation to non-agricultural species (Smith
and McClean, 1989; Brewer and Smith, 1995, 1997). Studies
with agricultural plant species have been performed only under
controlled conditions (Brewer and Smith, 1994; Ishibashi and
Terashima, 1995; Hanba et al., 2004).
Liu and Kang (2006a) reported the work of Yang et al.
(2000), who found increases in wheat (Triticum aestivum
L.) photosynthesis in sprinkler irrigated areas. Maize and
alfalfa are among the main sprinkler irrigated field crops
worldwide, but there is no information available about the
effect of daytime sprinkler irrigation on their photosynthesis.
The objective of this study was to determine whether the net
photosynthesis of maize and alfalfa is affected by sprinkler
irrigation in a solid-set system during daytime irrigation events
and to determine what mechanisms might affect it.
Experimental Site
Field experiments were conducted in Zaragoza, Spain
(41°43´ N, 0°48´ W, 225 m asl). Maize was grown in a 2.34-ha
field during 2009 and 2010 (Exp. 1) and alfalfa was grown in a
2.0-ha field during 2009, 2010, and 2011 (Exp. 2). The distance
between the two experiments was about 1 km. Both of them
were located in the middle of an irrigated 8000-ha area where
the main crops are maize, alfalfa, and wheat. The minimum
distance of the experimental plots to the edge of the irrigated
area was 2.5 km. The soil was clay loam in Exp. 1 and sandy
loam in Exp. 2. Both soils are classified as Typic Xerofluvent
(Table 1). The climate in the area is Mediterranean semiarid. In
the last 10 yr (2003–2012) annual daily mean air temperature
was 14.1°C, annual total precipitation was 298 mm, and annual
reference evapotranspiration (ETo) was 1243 mm.
Experimental Layout
A solid set sprinkler irrigation system was installed in both
experimental fields (Fig. 1 and 2). The sprinkler spacing was
square, 18 by 18 m in Exp. 1, and 15 by 15 m in Exp. 2. The
impact sprinkler and nozzles were manufactured in brass
(RC-130, Riegos Costa, Lérida, Spain). The sprinkler has a
vertical throw angle of 25°, the nozzle diameters were 4.4 mm
(main) and 2.4 mm (auxiliary), and the nozzle height was
2.5 m (Exp. 1) and 2.2 m (Exp. 2) above the soil surface. The
nozzle operating pressure was kept around 0.3 MPa with a
hydraulic pressure control valve. Sprinkler application rates
were 5 mm h–1 (Exp. 1) and 7.5 mm h–1 (Exp. 2). The irrigation
volume was measured with an electromagnetic flow meter
(Promag 50, Endress+Hauser, Reinach, Switzerland) which has
a measurement error of ±0.5%. The meteorological conditions
during the irrigation events were measured with an automated
weather station (thereafter named as the weather grass station)
located at the experimental farm, at 1 km from Exp. 1 and
50 m from Exp. 2. The weather grass station is located over
grass following the reference conditions defined by Allen et al.
(1998).
The field of Exp. 1 had been grown with maize the previous
year and the crop residues incorporated to the soil with
conventional tillage. Maize cultivar Pioneer PR34N43
(comparative relative maturity 110) was planted on 21 Apr.
2009 and 20 Apr. 2010 (Exp. 1), in rows 0.75 m apart at a

1516 Agronomy Journal  •  Volume 105, Issue 6  •  2013


Fig. 1. General layout of Exp. 1. Twelve irrigation sectors
irrigated independently by four sprinklers each. The irrigation
sectors with even numbers were irrigated during daytime in
2009 and during nighttime in 2010. The irrigation sectors with
odd numbers were irrigated during nighttime in 2009 and
during daytime in 2010. Irrigation was characterized (water
losses and irrigation uniformity) in sectors 5 and 6. The arrow
indicates where is the north (N).
planting density of 87,000 seeds ha–1. Fertilization consisted
of 50 kg ha–1 N, 100 kg ha–1 P2O5 and 100 kg ha–1 K 2O
applied preplant, and 200 kg ha–1 N applied with the
irrigation water split between the V6 and V12 growth stages
(Ritchie et al., 1996). Weeds were controlled by applying
acetochlor (2-chloro-N-(ethoxymethyl)-N-(2-ethyl-6-
methylphenyl)acetamide) and terbuthylazine (6-chloro-N-
(1,1-dimethylethyl)-N9-ethyl-1,3,5-triazine-2,4-diamine)
preemergence. Chlorpyrifos (O,O-diethyl O-3,5,6-trichloro-
2-pirydyl phosphorothioate) was applied at V14 through the
irrigation system for insect control. Alfalfa cultivar Aragon
was planted after conventional tillage at 35 kg ha–1 on March
2008 (Exp. 2). Fertilization consisted of 120 kg ha–1 P2O5 and
150 kg ha–1 K 2O applied each year in March. Six cuttings were
made every experimental year, starting on April and with an
approximated interval of 1 mo. No herbicides were necessary in
this crop but two to three insecticide applications were made
each year to control pests.
The field of Exp. 1 had 12 irrigation sectors (main) irrigated
by four sprinklers each (Fig. 1). The borders of the field were
irrigated independently of the main irrigation sectors. In this
experiment, maize was irrigated at nighttime until the crop
was well established (V6–V8 growth stage). Thereafter, two
irrigation time treatments were established (daytime and
nighttime), so six irrigation sectors were irrigated at daytime
and the other six at nighttime. Thus, at each daytime irrigation
event, microclimatic and physiological measurements (see
later in the text) made in one of the six daytime irrigated
sectors could be compared with those made in one of the
six nighttime irrigated sectors (which was not irrigated at
that time). The irrigation sectors irrigated during daytime
in 2009 were irrigated during nighttime in 2010 (Fig. 1).
Daytime irrigation generally started at 1000 GMT while
nighttime irrigation started at 2200 GMT of the previous day.
The weekly irrigation amount was generally applied in two
Agronomy Journal  •  Volume 105, Issue 6  •  2013 1517
Fig. 2. General layout of Exp. 2. Two irrigation sectors (A
and B) not irrigated at the same time. The area within each
irrigation sector where irrigation was characterized (water
losses and irrigation uniformity) is shown (IE). The arrow
indicates where is the north (N).
irrigation events that lasted 4 to 6 h. The area surrounded by
the four sprinklers of each irrigation sector was considered for
the measurements (Fig. 1). The area outside was disregarded
because it received water from two different irrigation sectors.
The soil matric potential was measured with a granular matrix
sensor (Watermark, Irrometer Co., Riverside, CA) in the
two irrigation sectors (one daytime irrigated and the other
nighttime irrigated) where net photosynthesis was measured.
Eight sensors were installed in each of these two irrigation
sectors at the same positions within the square defined by the
four sprinklers: four within the plant rows (two at 0.2-m depth
and two at 0.6-m depth) and four at the center of the interrow
space (two at 0.2-m depth and two at 0.6-m depth). The eight
sensors in each irrigation sector were connected to a datalogger
(Microisis, Sistemes Electrónics Progres, Bellpuig, Spain),
which recorded the measurements each hour. A soil matric
potential threshold of –0.080 MPa, which corresponded
to a 30% soil water depletion, was used to indicate probable
water stress in the experiment (Urrego-Pereira et al., 2013a).
Complete details of the experiment can be found in Urrego-
Pereira et al. (2013a).
The field of Exp. 2 was divided in two irrigation sectors 1.0
ha each (Fig. 2). In this experiment, alfalfa was irrigated twice
or thrice per week at each irrigation sector A and B (Fig. 2).
The duration of the irrigation events was generally limited
to 3 h. The applied irrigation water depth was the same at
each irrigation sector, but irrigations were not simultaneous.
Generally, at the beginning and end of the week the irrigation
sector A was irrigated during daytime periods and the
irrigation sector B was irrigated during nighttime the following
day. By the middle of the week, the irrigation sector A was
irrigated during nighttime and the irrigation sector B during
daytime the same day. Thus, at each daytime irrigation event,
measurements made in the irrigation sector being irrigated
could be compared with those made in the other nighttime
irrigation sector, which was not irrigated at that time. Daytime
irrigations started generally at 1000 GMT, while nighttime
irrigations started at 100 GMT.
 Crop water requirements were computed following the FAO
approach (Allen et al., 1998). Reference ETo was computed
with the FAO Penman–Monteith method from meteorological
data obtained from the weather grass station. Crop coefficients
(Kc) were calculated in the case of maize as a function of
thermal time developed in the same location (Martínez-Cob,
2008) or in the case of alfalfa from tabulated values (Allen et
al., 1998) locally adjusted by García-Vera and Martínez-Cob
(2004). Daily maize and alfalfa evapotranspiration (ETc) was
calculated from the corresponding daily values of ETo and Kc.
Next, the crop irrigation requirements were calculated weekly
as the difference between ETc and the effective precipitation,
which was estimated as 75% of precipitation (Dastane, 1978).
The irrigation amount applied to the crop was equal to the
crop irrigation requirements plus the measured water losses.
To evaluate the water losses of the irrigation events, a grid of
25 catch cans separated at 3.6 (Exp. 1) or 3.0 m (Exp. 2) was
installed within the square area delimited by four sprinklers
in two irrigation sectors (one daytime irrigated and the other
nighttime irrigated) of Exp. 1 and in the irrigation sectors A
and B of Exp. 2. The catch cans were made of plastic, had a
diameter of 0.18 m and were located just above the crop canopy.
After each irrigation event, the water amount collected in each
catch can was measured. WDEL was calculated as:
I fm − I cc
WDEL = 100
I fm
where Ifm, applied irrigation amount calculated from irrigation
water measured with the flow meter; Icc, mean irrigation
amount measured at the 25 catch cans. The coefficient of
uniformity of the irrigation (CU) was computed from
the water amount collected on the catch cans using the
methodology described by Christiansen (1942).
Microclimatic and Canopy Temperature
Changes Due to Irrigation
In Exp. 1 an automatic weather station was installed in the
center of one daytime irrigated sector and a second station in
one nighttime irrigated sector (Fig. 1). In Exp. 2 an automatic
weather station was installed in the center of each of the
two irrigation sectors (Fig. 2). These four weather stations
were used for continuously recording the air temperature
and VPD during the crop season so comparisons could be
made between the sector that was irrigated and the sector
that was not irrigated in each of the experiments during the
irrigation events. One temperature and relative humidity
probe (HMP45C, Vaisala, Helsinki, Finland) was installed at
each weather station at 1.0 m above the crop canopy of maize
and at 2.0 m above the soil surface of the alfalfa experiment.
Each probe was installed inside a shield URS1 (Campbell
Scientific, Logan, UT) which protected it from irrigation
water and solar radiation. The accuracy of the probes was
±0.3°C for temperature and ±3% for relative humidity. The
air temperature and relative humidity were measured every 10
s and the 5-min mean values were computed and recorded in
a datalogger (CR10X, Campbell Scientific Inc, Logan, UT).
The VPD was calculated from the air temperature and relative
humidity data (Allen et al., 1998).
1518 Agronomy Journal  •  Volume 105, Issue 6  •  2013
An infrared thermometer (Apogee Instruments Inc.,
Roseville, CA) was installed in each of the automatic weather
stations to measure the crop canopy temperature. The infrared
thermometer was located at 1.0 m above the crop canopy with
an angle of 45° and was oriented towards the north. The model
IRR-P, with an accuracy of ±0.1°C, was used in Exp. 1, and
the model IRTS-P, with an accuracy of ±0.3°C, was used in
Exp. 2. Canopy temperature measurements were taken at 10 s
intervals and average values every 5 min were recorded in the
dataloggers. Due to a technical problem canopy temperature
data from 2010 were lost.
Leaf Wettability and Surface Topography
Leaf wettability was determined by measuring advancing
contact angles of drops of double-distilled water at 25°C using
a Drop Shape Analysis System (DSA100, Krüss, Hamburg,
Germany). Contact angles were measured on intact adaxial and
abaxial maize and alfalfa leaf surfaces (10 replications). Two
microliter water drops were deposited on to the surfaces with a
manual dosing system holding a 3-mL syringe (0.5 mm diam.
needle). Side view images of the drops were captured at a rate of
six frames s–1. Contact angles were automatically calculated by
fitting the captured drop shape to the one calculated from the
Young–Laplace equation (Khayet and Fernández, 2012).
On 9 Aug. 2010 four maize leaves (ear leaf) were taken from
plants of the irrigated and not irrigated sectors at different
times (before the irrigation started, during the irrigation at 2,
3, and 4 h after the irrigation started, and at 1 and 3 h after the
irrigation finished). Water contact angles were measured on
five portions of each leaf. On 12 July 2010 eight plant samples
were taken from the irrigated and not irrigated sectors of
alfalfa just after the irrigation finished and the water contact
angles were measured in five leaves of each plant. The surface
topography of adaxial and abaxial gold sputtered, intact maize
and alfalfa leaf surfaces was examined with a scanning electron
microscope (SEM) (S-3400 N, Hitachi, Tokyo, Japan) (Khayet
and Fernández, 2012).
Net Photosynthesis
At each measurement day, net photosynthesis was measured
simultaneously in one sector being irrigated and another
one not irrigated at the same time. Data were collected from
1 h before the irrigation started until 2 h after the irrigation
finished, so data were collected for 6 to 9 h. For this task all
the equipment (chamber, pump, gas analyzer, flowmeter,
thermocouples) was replicated, so two sets of equipment were
used (one in the irrigated sector and another one in the not
irrigated sector).
Two automated transient-state closed-system canopy
chambers for CO2 exchange determination were used.
The chamber (Tecno El, Formello, Italy) is similar to that
described by Steduto et al. (2002). It had a main module that
is a rectangular box with five transparent polycarbonate walls
(1.5 mm thick), held together by a narrow aluminum angular
frame. The chamber was open in the bottom, had a ground
surface area of 0.75 m2 (1.0 by 0.75 m), and had a height of
0.5 m. It has a metal base that is inserted 5 cm into the soil. For
the maize crop, other modules with the same ground surface
area but with a height of 1.0 m and open in the base and in the
Table 2. Characteristics of the irrigation events and mean meteorological conditions during the irrigation events measured at the
weather grass station. The mean photosynthetically active radiation (PAR) reaching the crop canopy during each irrigation event is
also shown.
Experiment and Irrigation Irrigation Air Wind
irrigation date duration applied WDEL† CU‡ temp. Air VPD§ speed PAR
hh:mm mm –––––––––– % –––––––––– °C kPa m s–1 µmol m–2 s–1
Exp. 1: Maize
9 July 2009 4:00 21 18.8 87 25.4 2.0 2.3 1810
16 July 2009 5:15 27 22.1 86 31.6 2.5 2.6 1694
23 July 2009 5:00 25 24.2 72 29.7 2.5 3.3 1726
27 July 2009 5:00 26 25.6 68 31.1 2.9 4.1 1406
30 July 2009 5:30 28 21.9 82 28.9 2.4 3.3 1651
24 June 2010 5:30 27 10.9 90 29.7 3.1 1.2 1818
28 June 2010 4:30 22 9.5 84 26.9 1.8 2.3 1554
20 July 2010 5:30 28 8.5 83 30.7 1.9 2.2 1544
2 Aug. 2010 4:00 20 21.6 67 27.0 1.5 3.4 1624
9 Aug. 2010 4:00 20 11.2 86 30.8 2.3 2.9 1310
Exp. 2: Alfalfa
29 Sept. 2009 3:00 22 0.8 85 22.8 1.3 1.2 1257
1 Oct. 2009 3:00 22 22.4 82 25.3 1.6 3.4 1163
5 Oct. 2009 3:00 22 5.7 81 24.7 1.2 1.7 1243
8 Oct. 2009 3:00 22 10.5 78 24.9 1.6 2.0 1278
12 July 2010 3:00 22 9.6 84 32.0 2.9 1.3 1520
16 June 2011 3:00 22 18.6 87 27.9 2.2 3.2 1737
21 June 2011 3:00 24 7.3 85 30.3 2.4 0.9 2069
23 June 2011 3:00 23 15.6 89 23.8 1.5 3.2 1929
18 July 2011 3:00 22 15.4 87 22.0 1.5 2.6 2055
20 July 2011 3:00 23 30.3 77 22.5 1.5 5.2 1985
4 Aug. 2011 2:45 20 11.5 88 29.0 2.2 1.3 1959
19 Sep. 2011 3:00 22 35.3 76 20.3 1.3 7.0 1584
21 Sep. 2011 3:00 22 10.4 86 23.9 1.3 1.1 1629
28 Sep. 2011 2:30 18 13.5 84 25.7 1.7 2.6 1410
† WDEL, wind drift and evaporation losses.
‡ CU, coefficient of uniformity of Christiansen.
§ VPD, vapor pressure deficit.

top were added as the crop grew. Thus, the canopy chamber at
the maximum height of maize was composed by two modules
of 1.0 m height each and on top the main module of 0.5 m
height. In the case of the alfalfa crop only the main module
of 0.5 m height was used. In the case of maize, the chamber
was located centered in a plant row, so it covered six plants in
a 1.0 m length portion of the plant row. Each module has four
fans (Ebmpapst, Mulfingen, Germany) mounted in the corners
which provide a total flux of 1.4 m3 min–1. The chamber
top-cover has a hinge on one side, is usually open but can be
moved to close the chamber to measure the CO2 exchange. The
top cover was maintained opened at an angle of 75°, so some
interception of the irrigation water occurred. The chamber
was in place only during the measurement days in order not to
interfere with the crop.
Two thermocouples (Campbell Sci., TCBR-3, Shepshed,
UK) not shielded were installed to measure the air temperature
at a 0.5 s interval at each chamber at half of its height, one was
inside the chamber while the other was close but outside of
the chamber. The photosynthetically active radiation (PAR)
incoming above the crop canopy was measured at an interval of
60 s (Delta-T, model BF3, Wynster, UK).
A miniature diaphragm pump (model 15D1150, GAST,
Benton Harbor, MI) was used to continuously extract the air
from inside of the chamber (from two positions in maize: 1.5
and 2.3 m above the soil surface; from one position in alfalfa:
0.3 m above the soil surface) and to conduct it to an infrared
gas analyzer (IRGA, model LI-7000, Li-Cor Inc., Lincoln,
NE). A flowmeter (Dwyer, model VFB-66-SSV-BFP, Michigan
City, IN) was used to get a constant flow of 5 L min–1 through
the IRGA system. After the air has passed through the IRGA it
was recirculated to the chamber. The IRGA was set to measure
the CO2 concentration of the air every 0.5 s.
The chamber top-cover was kept open, except for a 50 s
period every 15 min. During the time that the chamber was
closed the four fans were stirring the air in each chamber
module. The net photosynthesis was calculated as the CO2
flux during the period that the top-cover was closed and
was determined with the concentration regression method
(Reicosky et al., 1990). We used a lag time of 10 s and a
calculation window of 20 s (40 values at a 0.5 s interval). Net
photosynthesis of maize was measured on 5 d on 2009 and 7 d
on 2010. Measurements were done after maize tasseling except
in the two dates of June 2010. Net photosynthesis of alfalfa was
measured on 4 d in 2009, 5 d in 2010, and 12 d days in 2011.
Maize and Alfalfa Yield
In Exp. 1, each of the 12 irrigation sectors (18 by 18m) was
harvested on 6 Oct. 2009 and 4 Oct. 2010 with a combine and
the grain weighed with a 1-kg precision scale. A subsample of

Agronomy Journal  •  Volume 105, Issue 6  •  2013 1519

grain was collected from each irrigation sector to measure the
grain moisture, a measurement used to adjust the grain yield to
a standard 140 g kg–1 moisture content. In Exp. 2, six samples
of the alfalfa aboveground biomass (0.75 m2) were taken at
each sector at each of the six alfalfa cuttings made per year. The
alfalfa samples were dried at 60°C.

Data Analysis
For each measurement day the net photosynthesis data set
was divided into four periods: (i) before the irrigation (before),
(ii) during the irrigation (during), (iii) 1 h after the irrigation
(1 h after), and (iv) 2 h after the irrigation (2 h after). For
each measurement day and period, the 15-min interval data
of net photosynthesis of the two treatments (irrigated and
not irrigated) were compared with a paired t test at a level of
significance of P ≤ 0.05. The measurement days when the net
photosynthesis before the irrigation was different between
the irrigated and not irrigated treatments were rejected.
Other variables measured at the same time in the irrigated
and not irrigated plots (air temperature and VPD, canopy
temperature, air temperature inside and outside the automated
canopy chamber) were analyzed also with a paired t test. Leaf
wettability in the irrigated and not irrigated plots was compared
with a t test. The daily soil matric potential in the daytime and
nighttime maize irrigated plots was compared with a paired
t test. The effect of irrigation time on maize yield was analyzed
with ANOVA. The Statgraphics 5.0 software was used to
analyze the data.
RESULTS
Characteristics of the Irrigation Events
The duration of irrigation events ranged from 4.0 to 5.5 h
for maize and from 2.5 to 3 h for alfalfa (Table 2). Taking into
account the sprinkler application rates for both experiments,
the average irrigation depths applied at each irrigation event
were similar for both species (maize: 24 mm, alfalfa: 22 mm).
For irrigation events of maize, the mean air temperature and
VPD ranged from 25 to 32°C and from 1.50 to 3.10 kPa,
respectively. For irrigation events of alfalfa, the mean air
temperature and VPD ranged from 20 to 32°C and from 1.20
to 2.90 kPa, respectively. The wind speed ranged from 1.2 to
4.1 m s–1 during the irrigation events of maize and from 0.9 to
7.0 m s–1 for those of alfalfa.
The WDEL during the irrigation events of maize ranged from
8 to 26% of applied water depth, while this range was wider
for irrigation events of alfalfa (1–35%) (Table 2). In general,
greater WDEL occurred with higher wind speed. The greatest
WDEL of maize (26%) and alfalfa (35%) were measured for
wind speeds of 4.1 and 7.0 m s–1, respectively. The CU values
for the irrigation events of maize ranged from 67 to 90% and for
the irrigation events of alfalfa from 76 to 89%. The lower CU
values were found in irrigation events with high wind speed. The
PAR ranged from 1310 to 1818 µmol m–2 s–1 during the maize
irrigation events, and from 1163 to 2069 µmol m–2 s–1 during
the alfalfa irrigation events.
The soil matric potential in Exp. 1 was similar in the daytime
and nighttime sprinkler irrigated maize plots and was always
higher than the water stress threshold for maize at this site (Fig. 3).
Fig. 3. Daily average values of soil matric potential for the
different irrigation time treatments in Exp. 1 (maize). Each
value is the average from eight probes: four probes installed
within the plant row (two at 0.2-m depth and another two
at 0.6-m depth), and another four probes installed halfway
between two plant rows (two at 0.2-m depth and another
two at 0.6-m depth). The dashed line indicates the soil matric
potential that can cause water stress in maize for the soil at
this experiment location. The stars indicate the days when
maize net photosynthesis was measured.
Microclimatic and Canopy Temperature
Changes Due to Irrigation
Reductions of air temperature and VPD and canopy
temperature during irrigation were observed as soon as
sprinkler irrigation started (Fig. 4 and 5). There was a
significant reduction of the air temperature measured above the
crop canopy on the irrigated treatment around 1.5°C (maize)
and 1.7°C (alfalfa) (Table 3). For both crops these significant
reductions lasted for 1 h after irrigation with an average
reduction of 0.6°C. Two hours after the irrigation finished
the air temperature values of the irrigated treatment matched
the air temperature values in the not irrigated treatment.
Likewise, air temperature measured with thermocouples inside
the canopy chamber was significantly lower at the irrigated
treatment as compared to the not irrigated treatment (on
average 3.8°C for maize and 3.5°C for alfalfa). This lower air
temperature determined in the irrigated treatment lasted
for 1 h after the irrigation finished and accounted for 1.9°C
(maize) and 1.3°C (alfalfa). Similar differences between the
irrigated and not irrigated treatments were observed for
the air temperature measured with thermocouples outside
the canopy chamber. The air temperature reductions due to
irrigation recorded with thermocouples were greater than those
reductions recorded with the Vaisala probes. This result could

1520 Agronomy Journal  •  Volume 105, Issue 6  •  2013

Fig. 4. Air temperature and vapor pressure deficit (VPD),
maize canopy temperature and net photosynthesis of
maize at 15-min interval from 1 h before until 2 h after the
end of the irrigation event on 30 July 2009 for the irrigated
and not irrigated treatments (y axis at the left side). The
photosynthetically active radiation (PAR) above the crop
canopy is shown in the y axis at the right side.
Fig. 5. Air temperature and vapor pressure deficit (VPD),
alfalfa canopy temperature and net photosynthesis of alfalfa
at 15-min intervals from 1 h before irrigation until 2 h after
the end of the irrigation event on 8 Oct. 2009 for the irrigated
and not irrigated treatments (y axis at the left side). The
photosynthetically active radiation (PAR) above the crop
canopy is shown in the y axis at the right side.

Table 3. Average air temperature and vapor pressure deficit (VPD) (measured above the crop canopy) and canopy temperature
of maize and alfalfa in the irrigated (Irrig) and not irrigated (Not irrig) plots during and after the irrigation events when photosyn-
thesis was measured (maize, 2009 and 2010; alfalfa, 2009, 2010, and 2011). Average air temperature measured with thermocouples
inside and outside the canopy chambers during the 50 s that the canopy chamber was closed is also provided.
During irrigation 1 h after irrigation 2 h after irrigation
Variable and crop N† Irrig Not irrig Irrig Not irrig Irrig Not irrig
Air temperature (°C)
Maize 10 27.6b‡ 29.1a 30.5b 31.1a 30.6b 30.7a
Alfalfa 14 23.6b 25.3a 26.8b 27.4a 27.4b 27.6a
VPD (kPa)
Maize 10 1.75b 2.19a 2.78b 2.91a 2.90a 2.90a
Alfalfa 14 1.30b 1.74a 2.05b 2.22a 2.29a 2.35a
Canopy temperature (°C)
Maize 5 24.4b 29.5a 28.0b 29.9a 29.0a 28.8a
Alfalfa 13 21.1b 27.0a 27.2b 28.2a 28.0a 27.1a
Temperature inside chamber (°C)
Maize 10 25.9b 29.7a 27.8b 29.7a 28.2a 28.7a
Alfalfa 14 23.6b 27.1a 26.7b 28.0a 27.0a 27.3a
Temperature outside chamber (°C)
Maize 10 26.3b 28.7a 28.2b 28.9a 28.0a 27.9a
Alfalfa 14 22.4b 26.3a 26.8b 27.4a 26.7a 26.9a
† Number of irrigation events.
‡ For each variable, crop and period of measurement, the values followed by different letters are significantly different according to a paired t test at the 0.05 probability level.

Agronomy Journal  •  Volume 105, Issue 6  •  2013 1521

Fig. 6. Advancing contact angles of a distilled water drop with the adaxial leaf surface of (A) maize and (B) alfalfa. Surface
topography (x200) of the adaxial leaf surface of (C) maize and (D) alfalfa. Surface topography (x200) of the abaxial leaf surface of
(E) maize and (F) alfalfa.

be related to the fact that thermocouples provided readings
inside the crop canopy while Vaisala probes provided readings
above the crop canopy. Cavero et al. (2009) found that the
microclimatic changes due to sprinkler irrigation are greater as
the measurement height over the soil surface decreases.
Sprinkler irrigation significantly reduced the air VPD
during irrigation by 0.44 kPa for both crops (Table 3). During
the first hour after the irrigation finished the significant VPD
reductions were 0.13 kPa (maize) and 0.17 kPa (alfalfa). For
both crops, the VPD recorded at the irrigated treatment
matched the recorded values at the not irrigated treatment
at the second hour after irrigation finished as in the case of
reductions observed for air temperature.
Sprinkler irrigation decreased maize canopy temperature
by 5.1°C during the irrigation event (Table 3). During the
first hour after the irrigation finished this decrease although
lower (1.9°C in average), was significant. The alfalfa canopy
temperature was also significantly reduced by 5.9°C during
sprinkler irrigation and 1 h after irrigation this significant
decrease was 1.0°C.
Leaf Wettability and Surface Topography
An example of the advancing contact angles of a distilled
water drop with the adaxial maize and alfalfa leaf surfaces
is provided in Fig. 6A and 6B. For maize, no significant
differences in water contact angles were determined in leaves
collected from the irrigated and not irrigated treatments in any
period of measurement (Fig. 7). For this species, lower water
contact angle values were found in the abaxial as compared
with the adaxial leaf surface. These water contact angles were
lower than 90° indicating that the leaves of maize are wettable.
Similarly, no water contact angle differences were found
between the irrigated and not irrigated treatments over the
adaxial and abaxial leaf surfaces of alfalfa (Fig. 8). However,
the contact angles of water drops with alfalfa leaf surfaces
were higher (121° for the adaxial surface and 125° for the
abaxial surface), which indicates that such surfaces are more
hydrophobic than maize leaves.
Examination of adaxial and abaxial leaf surfaces by
SEM provided evidence that the leaves of both species are
amphistomatous (Fig. 6C, 6D, 6E, 6F). The maize leaf surface
(Fig. 6C, 6E) was found to be more flat than the alfalfa leaf
surface, which had a more rough topography associated with
the shape of epidermal cells (Fig. 6D, 6F).
Net Photosynthesis
During the 50-s period that the canopy chamber was closed,
there was a slight increase of the air temperature inside the
canopy chamber as compared to the air temperature measured
outside the canopy chamber. This increase occurred in both the

1522 Agronomy Journal  •  Volume 105, Issue 6  •  2013

 Fig. 8. Contact angle of water droplets (θ) with the adaxial and
abaxial alfalfa leaf surfaces of the irrigated and not irrigated
treatments on 12 July 2010. Values are the means of eight
plants (five leaves from each plant were measured) and error
bars are the standard errors. For each type of leaf surface
no differences were found between the irrigated and not
irrigated treatments after a t test at P ≤ 0.05.

first measurement made) the net photosynthesis of maize at the

Fig. 7. Contact angle of water droplets (θ) with the adaxial and
abaxial maize leaf surfaces of the irrigated and not irrigated
treatments on 9 Aug. 2010. Values are the means of four
leaves from different plants and error bars are the standard
errors. For each time and type of leaf surface no differences
were found between the irrigated and not irrigated
treatments after a t test at P ≤ 0.05.
irrigated and not irrigated treatments and in average was below
1 and 1.2°C for maize and alfalfa, respectively (Table 3).
The time evolution of the net photosynthesis of maize
recorded at 15-min intervals on 30 July 2009 at the irrigated
and not irrigated treatments is shown in Fig. 4. The net
photosynthesis of maize was similar for both treatments before
irrigation. However, 15 min after the irrigation started (the
irrigated treatment decreased compared to the not irrigated
treatment. This lower photosynthesis measured in the irrigated
treatment remained until the end of irrigation. As soon as
irrigation finished, the net photosynthesis of irrigated maize
plants rose to the levels of not irrigated plants.
Two of the measurement days for net photosynthesis of
maize in 2010 were rejected from the analysis because there
were significant differences between the two treatments before
irrigation started. Thus, 10 irrigation events were available to study
the effect of sprinkler irrigation on maize net photosynthesis
(Table 4). Sprinkler irrigation significantly reduced the maize
net photosynthesis on 8 of the 10 irrigation events during the
irrigation period. In the other two irrigation events, there were
no differences between the two treatments. The reduction of
net photosynthesis of maize at the irrigated treatment ranged
from 10 (16 July 2009) to 41% (27 July 2009). Considering
all the irrigation events, a mean 19% reduction of maize net
photosynthesis was found in the irrigated treatment compared to
the not irrigated treatment during the irrigation event.

Table 4. Net photosynthesis of maize in the irrigated (Irrig) and not irrigated (Not irrig) plots before, during, and after the irriga-
tion events.
Net photosynthesis
Before irrigation During irrigation 1 h after irrigation 2 h after irrigation
Irrigation date Crop height Irrig Not irrig Irrig Not irrig Irrig Not irrig Irrig Not irrig
m —————————————————————— µmol m–2 s–1 —————————————————————
9 July 2009 2.40 47.8†a 56.5a 58.0b 70.3a 48.5a 69.3a 60.5b 68.8a
16 July 2009 2.40 82.3a 76.2a 76.4b 85.0a 49.5a 61.3a 20.8a 15.0a
23 July 2009 2.40 30.4a 35.8a 51.1b 68.8a 61.1a 66.8a – –
27 July 2009 2.40 73.9a 76.2a 39.0b 66.4a 21.7b 35.9a 9.8a 19.4a
30 July 2009 2.40 53.5a 55.6a 52.5b 66.7a 39.5a 42.6a 16.6a 17.2a
24 June 2010 1.05 40.0a 44.5a 39.7b 47.5a 38.5a 34.7a 23.5a 20.8a
28 June 2010 1.40 55.7a 53.9a 62.6a 63.6a 61.3a 59.0a 47.9a 52.9a
20 July 2010 2.45 61.2a 66.9a 65.4b 82.5a 39.1a 42.0a 13.1a 22.8a
2 Aug. 2010 2.45 46.1a 51.8a 55.8b 81.8a 50.5b 72.0a 32.7b 51.3a
9 Aug. 2010 2.45 44.9a 46.8a 53.2a 52.6a 38.0a 14.5a – –
Mean 53.6 56.4 55.4 68.5 44.8 49.8 28.1 33.5
† For each irrigation date and period the values followed by different letters are significantly different according to a paired t test at the 0.05 probability level.

Agronomy Journal  •  Volume 105, Issue 6  •  2013 1523

Fig. 9. Relationship between the net photosynthesis of maize and alfalfa and the temperature inside of canopy chamber during
sprinkler irrigation at the irrigated and not irrigated treatments along the irrigation seasons in 2009 and 2010 (maize) and 2009,
2010, and 2011 (alfalfa). Each point is the average value of an irrigation date. The dotted line indicates the temperature threshold
when photosynthesis of maize decreases as temperature decreases (Duncan and Hesketh, 1968; Tollenaar, 1989; Labate et al.,
1991; Wolfe, 1991; Kim et al., 2007). PAR is the photosynthetically active radiation, and V8 is the eight leaves growth stage (Ritchie
et al., 1996).

During the first hour after the irrigation finished the why no significant net photosynthesis differences were found
net photosynthesis of maize at the irrigated treatment was between the irrigated and not irrigated maize treatments in
significantly lower as compared to the not irrigated treatment this irrigation event.
in 2 d with a reduction of 30 to 39%. In the other 8 d there The time evolution of the net photosynthesis of alfalfa
were not differences between the two treatments (Table 4). recorded at each 15-min interval on 8 Oct. 2009 at the
Considering all the irrigation events, during the first hour after irrigated and not irrigated treatments is shown in Fig. 5.
the irrigation finished the maize net photosynthesis was 10% Before irrigation started the net photosynthesis of alfalfa was
lower in the irrigated treatment compared to the not irrigated similar for both treatments. Around 1 h after the onset of the
treatment. Finally, during the second hour after irrigation irrigation, the net photosynthesis of alfalfa was slightly higher
of maize finished the net photosynthesis of the irrigated than that of not irrigated plants. Once the irrigation finished
treatment was lower as compared to the not irrigated treatment the net photosynthesis of alfalfa at the irrigated treatment
in 2 d with a reduction of 12 to 36%. In the other 6 d, there began to be similar to that of the not irrigated treatment
were not differences between the two treatments. Considering and in the second hour after the irrigation finished the net
all the irrigation events, during the second hour after the photosynthesis of alfalfa was similar in the two treatments.
irrigation finished the maize net photosynthesis was 16% Four of the measurement days for net photosynthesis of
lower in the irrigated treatment compared to the not irrigated alfalfa in 2010 and three in 2011 were discarded from the
treatment.
The lower net photosynthesis of maize during the irrigation
event at the irrigated treatment was related with the lower
air temperatures inside the canopy chamber reached at this
treatment compared to the not irrigated treatment (Fig. 9).
Thus, the air temperature at the irrigated treatment was mostly
below the optimum range of 27 to 35°C (Duncan and Hesketh,
1968; Tollenaar, 1989; Wanjura and Upchurch, 2000; Crafts-
Brandner and Salvucci, 2002; Kim et al., 2007) and net
photosynthesis decreased as air temperature within the canopy
chamber decreased below 27°C.
Low values of net photosynthesis of maize at the not
irrigated treatment were found with incomplete canopy cover
(24 June 2010, maize height of 1.05 m) and when the PAR was
low (9 Aug. 2010, 1310 µmol m–2 s–1). In the latter date similar
values of net photosynthesis were measured at air temperatures
of 27°C (irrigated treatment) and 30°C (not irrigated Fig. 10. Relationship of the difference between net photo­
treatment). Figure 10 shows that during sprinkler irrigation synthesis of maize measured during irrigation in the irrigated
the decrease of maize net photosynthesis at the irrigated and not irrigated treatments and the photosynthetically
active radiation (PAR) along the irrigation seasons in 2009
treatment compared to the not irrigated treatment was less as and 2010. Each value corresponds to a 15-min interval
the PAR was lower. The low PAR on 9 Aug. 2010 may explain measurement.

1524 Agronomy Journal  •  Volume 105, Issue 6  •  2013

analysis because there were significant differences between the Maize and Alfalfa Yield
two treatments before onset of irrigation. Thus, 14 irrigation In the case of the Exp. 1 the yield of nighttime irrigated
events were available to assess the effect of sprinkler irrigation maize ranged from 15.0 Mg ha–1 (2009) to 16.5 Mg ha–1
on alfalfa net photosynthesis (Table 5). During the irrigation (2010). Daytime irrigation significantly (P < 0.05) decreased
period, sprinkler irrigation did not affect the alfalfa net the maize yield by 9% compared with nighttime irrigation
photosynthesis in half of the irrigation events, significantly in 2010. There was a 5% yield decrease for daytime irrigation
increased the net photosynthesis in 5 of the 14 monitored compared with nighttime irrigation in 2009 that was
irrigation events, and significantly decreased the alfalfa net significant at P = 0.10. Complete details about the effect of
photosynthesis in 2 of the 14 monitored irrigation events. irrigation time on the growth of maize can be found in Urrego-
The increase of net photosynthesis of alfalfa at the irrigated Pereira et al. (2013a). In Exp. 2, the alfalfa forage yield was
treatment ranged from 10% (21 June 2011) to 30% (16 June 24.2 Mg ha–1 in 2009, 20.5 Mg ha–1 in 2010 and 22.9 Mg ha–1
2011), while the decrease of net photosynthesis of alfalfa at the in 2011. Yield differences of alfalfa forage between the two
irrigated treatment ranged from 15% (23 June 2011) to 30% sectors ranged from 2 to 6%.
(28 Sept. 2011). Considering all the irrigation events, similar
net photosynthesis was found for alfalfa in the irrigated and DISCUSSION
not irrigated treatments during the irrigation event. The microclimatic changes (reduction of air temperature
During the first hour after the irrigation of alfalfa finished, and VPD) during and after sprinkler irrigation recorded in
the net photosynthesis at the irrigated treatment was higher this work were similar to previous reports (Thompson et al.,
as compared to the not irrigated treatment on 4 d with an 1993; Tolk et al., 1995; Saadia et al., 1996; Cavero et al., 2009),
increase of 14 to 57%. In the other 10 d, there were not and induced plant physiological responses such as the decrease
differences between the two treatments. Considering all the in canopy temperature (Steiner et al., 1983; Tolk et al., 1995;
irrigation events, during the first hour after the irrigation of Saadia et al., 1996; Liu and Kang, 2006a,b; Cavero et al.,
alfalfa finished the net photosynthesis was 12% higher in the 2009). The decrease in canopy temperature of maize found in
irrigated treatment as compared to the not irrigated treatment. our study was similar to that found by Tolk et al. (1995) using
Finally, during the second hour after stopping the irrigation a lateral-move sprinkler irrigation system and by Cavero et al.
of alfalfa, the net photosynthesis at the irrigated treatment (2009) using a solid-set sprinkler irrigation system. The slightly
was higher as compared to the not irrigated treatment in 3 d, lower mean decrease of canopy temperature of maize compared
with an increase of 25 to 55%. In the other 11 d, there were to alfalfa due to sprinkler irrigation was probably due to a
not differences between the two treatments. Considering all lower water application rate for maize (5 mm h–1) as compared
the irrigation events, during the second hour after irrigation of with alfalfa (7.5 mm h–1). A higher application rate can have a
alfalfa finished the net photosynthesis was 13% higher in the higher cooling effect on crops (Cavero et al., 2009).
irrigated treatment compared to the not irrigated treatment. It has been reported that sprinkler irrigation increases leaf
In the case of the alfalfa crop, although the temperature inside water potential of maize (Cavero et al., 2009). This plant
the canopy chamber decreased due to sprinkler irrigation, no physiological change should increase maize photosynthesis
relationship was found between the net photosynthesis and the according to the work of Boyer (1970a,1970b) and Beadle
air temperature inside the canopy chamber within the range of et al. (1973). Thus, increased leaf water potential due to mist
temperature measured (20–32°C) (Fig. 9). irrigation resulted in a yield increase for southern peas (Howell
Table 5. Net photosynthesis of alfalfa in the irrigated (Irrig) and not irrigated (Not irrig) plots before, during and after the irriga-
tion events.
Net photosynthesis
Before irrigation During irrigation 1 h after irrigation 2 h after irrigation
Irrigation date Crop height Irrig Not irrig Irrig Not irrig Irrig Not irrig Irrig Not irrig
m ————————————————————— µmol m–2 s–1 —————————————————————
29 Sept. 2009 0.20 17.9†a 19.8a 20.5a 19.9a 19.3a 20.5a 12.4a 13.7a
1 Oct. 2009 0.25 25.2a 22.1a 25.8a 20.8b 23.2a 14.8b 17.5a 11.3b
5 Oct. 2009 0.35 26.5a 27.7a 29.3a 29.3a 24.1a 21.1b 23.7a 18.9b
8 Oct. 2009 0.42 26.1a 26.5a 30.3a 27.2b 21.2a 18.8a 14.9a 11.4a
12 July 2010 0.50 34.8a 36.0a 29.9a 24.0b 30.0a 23.3a 26.2a 20.3b
16 June 2011 0.35 26.9a 26.4a 27.8a 21.4b 22.9a 16.9b 17.1a 13.7a
21 June 2011 0.40 30.5a 28.7a 31.3a 28.4b 29.5a 28.6a 3.6a 4.8a
23 June 2011 0.45 35.1a 37.8a 33.6b 39.4a 35.5a 35.9a 33.0a 34.1a
18 July 2011 0.30 26.1a 26.4a 30.4a 27.0a 29.7a 26.7a 26.0a 22.0a
20 July 2011 0.35 31.7a 26.9a 34.2a 30.0a 31.0a 27.9a 27.9a 24.4a
4 Aug. 2011 0.50 25.0a 26.7a 21.8a 25.6a 22.8a 22.6a 22.1a 19.6a
19 Sept. 2011 0.20 20.1a 20.6a 22.3a 21.8a 23.2a 20.8a 21.8a 18.5a
21 Sept. 2011 0.25 23.6a 22.9a 22.3a 23.1a 26.1a 19.7b 20.4a 19.0a
28 Sept. 2011 0.40 28.0a 23.5a 21.4b 30.8a 23.3a 25.4a 23.4a 25.1a
Mean 27.0 26.6 27.2 26.3 25.8 23.1 20.7 18.3
† For each irrigation date and period the values followed by different letters are significantly different according to a paired t test at the 0.05 probability level.

Agronomy Journal  •  Volume 105, Issue 6  •  2013 1525


Fig. 11. Leaves of maize (wettable) and of alfalfa (non-
wettable) during sprinkler irrigation.
et al., 1971). Liu and Kang (2006 a, 2006b) reported a higher
yield of sprinkler irrigated winter wheat as compared to surface
irrigated.
Our field experiments showed that sprinkler irrigation
clearly decreased the net photosynthesis of maize by an average
of 19% during the irrigation event and slightly decreased
the net photosynthesis of maize by an average of 13% during
the 2 h following the irrigation event. However, sprinkler
irrigation did not affect the net photosynthesis of alfalfa
during the irrigation event and slightly increased the net
photosynthesis of alfalfa by an average of 12% during the
2 h following the irrigation event. Soil matric potential data
indicate that the decreased net photosynthesis of maize during
daytime sprinkler irrigation could not be attributed to water
stress. There is one plant morphological trait and one plant
physiological trait that can explain the different response of net
photosynthesis to sprinkler irrigation of maize and alfalfa.
The plant morphological trait that may explain the different
response of net photosynthesis to sprinkler irrigation of maize
and alfalfa is leaf wettability (Smith and McClean, 1989;
Brewer and Smith, 1994, 1995; Ishibashi and Terashima, 1995;
Hanba et al., 2004). Measurement of advancing contact angles
of water drops showed that maize leaf surfaces are wettable
(water contact angles < 90°), while the alfalfa leaf surfaces are
more hydrophobic and non-wettable (water contact angles
>120°). It has been reported that the wettability of maize
leaves changes with phenology with the first five to six leaves
being non-wettable while older leaves are wettable (Bianchi
et al., 1985; Beattie and Marcell, 2002), as observed in our
work. Bradley et al. (2003) reported that some Medicago spp.
have non-wettable leaves. In a growth chamber study, Hanba
et al. (2004) reported that moistening the leaves of a wettable
species (bean, Phaseolus vulgaris L.) decreased photosynthesis,
while moistening the leaves of a non-wettable species (pea,
Pisum sativum L.) slightly increased photosynthesis. For the
maize crop (wettable species), the sprinkler irrigation water
drops covered a large portion of the leaf surface (Fig. 6A, 11)
and represented a barrier to CO2 diffusion into the mesophyll,
which led to reduced photosynthesis. Hanba et al. (2004)
found that the reduction of net photosynthesis on the wettable
species occurred shortly after moistening the leaves during 10
min and that the photosynthesis increased rapidly as the layer
of water evaporated. However, they found that the reduction of
net photosynthesis was irreversible when the leaves have been
exposed to moistening for a long period of time (72 h) due to a
16% decrease in stomatal conductance and a 55% reduction in
the amount of Rubisco (ribulose-1,5-biphosphate carboxylase).
In our work, we found that the net photosynthesis of maize
1526 Agronomy Journal  •  Volume 105, Issue 6  •  2013
was reduced by sprinkler irrigation shortly after the irrigation
started but given the short duration of irrigation (4–6 h)
the net photosynthesis of maize was only reduced in 2 out of
10 d along the first and second hour after irrigation finished.
Therefore, during sprinkler irrigation of maize the CO2
diffusion and fixation processes were limited by the irrigation
water drops covering the leaf surface, and photosynthesis was
not totally recovered along the 2 h following the irrigation.
In the case of the alfalfa crop (non-wettable species) no
water barrier to CO2 uptake was formed (Fig. 11), therefore
the CO2 diffusion and fixation processes were not affected
and the sprinkler irrigation had a slight positive effect for
photosynthesis probably because of the lower VPD (Smith and
McClean, 1989; Brewer et al., 1991; Brewer and Smith, 1995,
1997).
The potential uptake of water by the foliage of the wettable
bean leaf was also considered by Hanba et al. (2004) in
association with the reduced photosynthesis of moist leaf
surfaces. Water may be taken up by the foliage via stomata,
the cuticle, cuticular cracks, and modified epidermal cells
such as trichomes (Fernández and Eichert, 2009). The area
of contact between water drops deposited on to maize leaf
surfaces is much higher than on the more hydrophobic alfalfa
leaves. Hence, a larger leaf area will be covered by water drops
in the more wettable maize leaf surface, which may facilitate
the process of foliar water uptake. However, the actual
mechanisms of foliar penetration by water are currently not
fully understood, and will be affected by the chemistry and
physical structure of every particular leaf surface (Khayet
and Fernández, 2012). Hence, more detailed water uptake
experiments with maize leaves will be required to clarify
whether the photosynthesis reduction recorded for irrigated
maize may be also linked to the potential absorption of
irrigation water by the foliage.
The plant physiological trait that may explain the different
response of net photosynthesis to sprinkler irrigation of maize
and alfalfa is the response of net photosynthesis to temperature
of each species. Lower net photosynthesis of maize during
sprinkler irrigation was related with lower temperatures
reached at the irrigated treatment while the net photosynthesis
of alfalfa was not affected by the lower temperatures at the
irrigated treatment. This could be due to the fact that the C4
plant species (maize) have a higher temperature optimum for
photosynthesis than C3 species (alfalfa) (Berry and Björkman,
1980). For maize, the optimum temperature for photosynthesis
has been established around 27 to 35°C (Duncan and Hesketh,
1968; Tollenaar, 1989; Wanjura and Upchurch, 2000;
Crafts-Brandner and Salvucci, 2002; Kim et al., 2007) and a
reduction in photosynthesis has been found when temperature
decreases below 27°C (Duncan and Hesketh, 1968; Tollenaar,
1989; Labate et al., 1991; Wolfe, 1991; Kim et al., 2007).
For alfalfa, a wide range of temperature optimum for
photosynthesis has been established (20–30°C) (Chatterton
and Carlson, 1981; Brown and Radcliffe, 1986; Al-Hamdani
and Todd 1990; Ziska and Bunce, 1994). Therefore, for maize
the air temperature measured at the irrigated treatment
(between 23 and 28°C) was below the optimum range for 80%
of the irrigation events probably reducing net photosynthesis.
However, in the case of alfalfa, the air temperature at the
irrigated treatment (between 20 and 28°C) was within the
optimum range for most irrigation events, and consequently
did not affect net photosynthesis.
Results from our work and previous at the same location
(Cavero et al., 2008) have found that daytime sprinkler
irrigation of maize decreased the grain yield by 5 to 10%
compared to nighttime sprinkler irrigation. This yield decrease
was mainly caused by the lower irrigation uniformity of
daytime irrigation due to the higher daytime wind speed
(Urrego-Pereira et al., 2013a). However, the lower net
photosynthesis of maize during sprinkler irrigation found in
our study indicates that part of the maize yield decrease with
daytime sprinkler irrigation could be due to the reduced net
photosynthesis. Most sprinkler solid-set systems at our site
(18 by 18 m spaced, 5 mm h–1) must be run for 12 h wk–1 to
apply enough water for maize during the main growing period.
Considering a 19% reduction of net photosynthesis during the
irrigation event this will result in an average 3% decrease of
maize yield if sprinkler irrigation is performed during daytime.
Although this is a slight maize yield decrease, considering that
alfalfa photosynthesis has a slight positive response to sprinkler
irrigation, when daytime sprinkler irrigation is necessary due to
constraints of the irrigation system, alfalfa should be the crop
daytime irrigated.
In hotter environments than in our site, the decrease of
temperature due to sprinkler irrigation could be beneficial
for maize if the reduced temperature remains within the
optimal range for photosynthesis. However, the decreased CO2
absorption due to the leaf surface deposited water barrier could
also decrease maize photosynthesis. Under sprinkler moving
systems the time that the plants are moistened is generally
lower compared to solid-set systems although physiological
changes can have a similar duration (Urrego-Pereira et al.,
2013b). Further studies are needed to clarify the impact of
sprinkler irrigation on the photosynthesis of field crops under
different environments and sprinkler irrigation systems.
CONCLUSIONS
Sprinkler irrigation decreased the net photosynthesis of
maize on 80% of days during the irrigation event and the mean
reduction was 19%. However, sprinkler irrigation increased
the net photosynthesis of alfalfa during the irrigation event
on 36% of days, decreased it on 14% of days, and did not affect
it on half of the days, so on average the net photosynthesis of
alfalfa was not affected. In the 2 h after the sprinkler irrigation
finished, the net photosynthesis of maize was reduced on 20%
of days but the net photosynthesis of alfalfa was increased on
21% of days.
The reduction of net photosynthesis of maize during
sprinkler irrigation was related both with the high wettability
of maize leaves, which reduced the exchange of CO2 , and with
the reduction of canopy and air temperature, which resulted
in temperatures below the optimum for photosynthesis of
maize. The low wettability of the leaves of alfalfa prevented
sprinkler irrigation water from interfering with the CO2
uptake and therefore did not decrease net photosynthesis.
Moreover, sprinkler irrigation did not decrease air and canopy
temperature below the optimum range for photosynthesis of
Agronomy Journal  •  Volume 105, Issue 6  •  2013 1527
alfalfa. The decrease of air VPD due to sprinkler irrigation
induced a slight increase of alfalfa net photosynthesis.
Daytime sprinkler irrigation with solid set systems under
similar climatic conditions to the study site should be avoided
for maize due to decreased net photosynthesis and yield.
Daytime sprinkler irrigation can be used for alfalfa because it
did not decrease net photosynthesis but the consequences of
irrigation time on yield should be studied.
Acknowledgments
This work was supported by the projects AGL2007-66716-C03-01
and AGL2010-21681-C03-01 (Ministerio de Economía y
Competitividad) of Spanish Government. We thank C. Merino,
M. Izquierdo, J. Gaudó, J.M. Asín, R. Santolaria, V. Pérez, and P.
Paniagua for the technical assistance in the field. We also thank V. del
Rio for measurements of leaf wettability and surface topography. Y.F.
Urrego-Pereira had a FPI grant of the Spanish government.
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