European Journal of Agronomy 9 (1998) 21–34

Nitrogen fertilizer can increase dry matter, grain production

and radiation and water use efficiencies for durum wheat
under semi-arid conditions
K. Latiri-Souki a,*, S. Nortcliff b, D.W. Lawlor c
a INRGREF BP 10-2080, Ariana, Tunisia
b University of Reading, Department of Soil Science Whiteknights, PO Box 233, Reading, RG6 2DW, UK
c IACR-Rothamsted, Biochemistry and Physiology Department, Harpenden, AL5 2JQ, UK
Accepted 22 January 1998

Abstract

Responses of durum wheat (Triticum durum, cultivar Karim) to application of nitrogen fertilizer and irrigation have
been analysed in the semi-arid areas of Tunisia. The aim is to establish the value, for crop production, of applying N
fertilizer under non-irrigated conditions and analyse the mechanisms of responses in two soil types: a clay soil at
Cherfech and a sandy soil at Nabeul. Irrigation increased above-ground dry matter production substantially on the
sandy soil but not on the clay soil. Nitrogen application also stimulated dry matter production substantially in non-
irrigated (rainfed) as well as irrigated conditions. The stimulation was greater under drought conditions than well-
watered conditions: 37% compared to 27%, respectively. Positive effects of N were obtained on both soils over all
years. The effect was due to increased LAI, and maintenance of green crop duration. This increased total dry matter,
which resulted in improved efficiencies of radiation and water use. However, the nitrogen use efficiency decreased
with nitrogen application. The mechanisms of the effects and the impact on durum wheat production and its
improvement under semi-arid conditions are considered. © 1998 Elsevier Science B.V.

Keywords: Durum wheat (Triticum durum) irrigation; Nitrogen; Radiation use efficiency; Semi-arid

1. Introduction Amami, 1984; Evans, 1993). The supply of nutri-

Water is the main factor limiting growth and
grain filling of cereals under semi-arid conditions
(Fisher, 1981; Kassam, 1981; Smith and Harris,
1981). As a result of the highly variable amount
and timing of rainfall between years, cereal pro-
duction is extremely variable (Papy, 1979; El
* Corresponding author. Tel: +216 1 719 630; fax: +216 1
717 951.
ents, in a suitable form, is also a major factor
determining cereal production (van Duivenbooden
et al., 1996) with nitrogen being the most impor-
tant applied fertiliser (Sinclair and Horie, 1989;
Grindlay, 1997).
Water deficits affect many plant processes; organ
development is slowed, and their growth rate is
decreased, resulting in fewer tillers and fewer,
smaller leaves in the crop so decreasing leaf area
per unit ground area (LAI ) (Legg et al., 1979).

1161-0301/98/$19.00 © 1998 Elsevier Science B.V. All rights reserved.

PII S 11 6 1 -0 3 0 1 ( 9 8 ) 0 0 0 22 - 7
22 K. Latiri-Souki et al. / European Journal of Agronomy 9 (1998) 21–34
Also, fewer ears with fewer spikelets and grains
are formed, which adversely affects grain yield
(Lawlor et al., 1981; Giunta et al., 1993). Drought
restricts the availability of carbon assimilates from
photosynthesis for vegetative growt and grain fill-
ing due to smaller stomatal conductance, which
not only decreases photosynthesis by slowing the
rate of CO entry into the leaf, but also slows the
2
rate of transpiration and water loss per unit leaf
area. Hence, both total assimilate production and
water use are decreased by the combination of
smaller LAI ( less photosynthetic and evaporative
surface area) and greater resistance to carbon
dioxide uptake and water loss (Day et al., 1981).
Thus, water deficits decrease dry matter production
during vegetative and reproductive growth and
thereby decrease yield ( Tanner and Sinclair, 1983).
Generally, the effects of water deficit on water loss
are greater than the effects on photosynthesis and
dry matter accumulation, so that water use effi-
ciency ( WUE ) (ratio of dry matter produced to
water consumed) may increase under dry com-
pared to wet conditions although total crop pro-
duction is generally severely restricted (Ritchie,
1983). Water consumption by crops is dependent
on several factors, including the local atmospheric
conditions (principally humidity and wind speed ),
the crop temperature and the soil water status
(Monteith and Elston, 1993).
Dry matter production is a function of the total
light energy intercepted (Monteith, 1977), which
drives photosynthesis and of the total amount of
CO assimilated minus losses by respiration. All
2
of these processes are affected by the leaf area
index and stomatal conductance (Tardieu et al.,
1991), and there are complex interactions between
them. An increase in LAI may reduce soil evapora-
tion, thus offsetting the increased loss through the
plants: depending on the relative importance of
these processes, the water use efficiency may be
improved (Ritchie, 1983). In agriculture, under
dry conditions, relatively small changes in condi-
tions may be important in biomass and yield
production.
Nitrogen is one of the most important nutrients
applied as a fertilizer, responsible to a great extent
for the large yields obtained from high input
agriculture (Greenwood, 1982; van Keulen and
Seligman, 1987). The supply of nitrogen increases
total biomass production and increases yield and
yield components. N affects production through
different mechanisms. At a cellular level, N
increases the cell number and cell volume; at the
leaf level, it increases the photosynthetic rate and
efficiency (Lawlor et al., 1988). The effect of
fertilizer N is to increase the amounts of the plant’s
metabolic components (e.g. proteins) shown by an
increase in the percentage of nitrogen in the tissue
(Greenwood et al., 1991). Increases in crop growth
rate are largely produced through an increase in
leaf area index, and also by an increase in radiation
use efficiency (RUE, dry matter produced per unit
of either incident radiation or intercepted radia-
tion) (Brown et al., 1987, Lawlor, 1995). Water
use efficiency also increases as consequences of
increasingly variance and radiation interception.
There is a conflict between the effects of water
deficit and of additional N fertilizers in crop
production (Dann, 1969; Anderson, 1985;
Gonzales Ponce et al., 1993). To increase biomass,
a larger LAI and a longer duration for which LAI
is maintained are required: these can be obtained
by adding fertilizer N. Increased biomass generally,
but not necessarily, results in increased grain yield.
Under drought conditions, smaller LAI and
shorter crop duration are desirable to decrease
water transpiration by the crop (Shepherd et al.,
1986; Cooper et al., 1987). Thus, the benefits of
applying nitrogen under semi-arid conditions will
depend on the frequency and intensity of drought
and on the amounts and timing of N applications.
In West Asia and North Africa, including Tunisia,
cereals are grown under semi-arid climatic condi-
tions where the total rainfall is low and variable
(Papy, 1979; Kassam, 1981; Smith and Harris,
1981; El Amami, 1984; Cooper et al., 1987). In
the experimental study reported in this paper,
durum wheat is considered as it is the most impor-
tant and widely grown crop in semi-arid areas in
Tunisia (Latiri-Souki et al., 1992). The application
of nitrogen fertiliser to crops grown in the semi-
arid conditions varies from year to year in Tunisia
(Aubry et al., 1994) as well as in other dry areas
(van Duivenbooden et al., 1996). In Tunisia, nitro-
gen is not always applied by farmers and when
applied, is generally applied late, after the plant’s
 K. Latiri-Souki et al. / European Journal of Agronomy 9 (1998) 21–34
apical differentiation to form ears is complete
(stage 4 or 5 in Feekes’ scale) and then only if the
early growing season has had adequate rain.
Although N fertilizers are subsidised in Tunisia,
they are not always applied by farmers because of
the extra costs involved and also because it is
commonly thought that application of N may
result in a lower yield if there is little rain.
However, there is inadequate evidence for this
supposition.
In this study, the combined effects of N and
water supply were measured to examine whether
application of N can be beneficial for durum
wheat, grown in a semi-arid area where water
deficits, sufficient to severely impair crop pro-
duction, are frequent. As responses to N and
irrigation vary with soil and weather conditions
( Thorne et al., 1988), they were tested on two soil
types with very different soil water holding capacity
and in different growing seasons. The questions
we test are: the positive effects of nitrogen fertilizer
on crop production obtained under well-watered
conditions also observed under dry conditions?
What are the mechanisms of the crop response to
nitrogen fertilizer under water deficit? Are results
dependent on climatic and soil conditions?
2. Materials and methods
2.1. Locations, climate, soil and water supply
Crops were grown at two locations and in three
different years in four field experiments ( Table 1)
with a variety of durum wheat (Triticum durum L
Table 1
Experiments and experimental treatments
Location Cherfech
Year 89–90
Abreviation L1Y1
Irrigation 2 levels
Nitrogen 2 levels
Treatments 4 4
Blocks 4 4
Subplots 16
Size of subplots 6×19 m
23
cultivar Karim) widely used in Tunisia.
Experiments were carried out at Cherfech in the
Mejerda valley (this site is designated L1,
Longitude: 10°3∞30◊E, latitude: 36°57∞30◊N, alti-
tude: +10.5 m) and Nabeul in the Cap Bon penin-
sula (designated L3, Longitude: 10°40∞15◊E,
latitude: 36°27∞30◊N, altitude: +24 m) in three
different growing seasons (1989–1990: designated
Y1; Y2: 1990–1991; Y3: 1991–1992).
The mean annual rainfall is 478 mm at Cherfech
and 450 mm at Nabeul, whereas the pan evapora-
tion varies from about 1.4 (January) to
8.4 mm day-1 (July) at Cherfech and 1.5 (January)
to 7.9 mm day− 1 (July) at Nabeul. The average
daily temperature at both locations is about 11°C
in January and 26°C in August.
The soil at Cherfech ( Table 2) is a silty clay
loam, organic matter content 2.2%, of high water
holding capacity. The water table is at an average
depth of 1.5 m ( UNDP-UNESCO, 1970). At
Nabeul ( Table 2), the soil is a very sandy and
homogeneous fluvisol of low (1.3%) organic matter
content (Chaabouni, 1983), with a small water
holding capacity and a water table below a depth
of 2 m.
2.2. Experimental design and experimental
treatments
2.2.1. Experimental design
The experimental design was a randomised split
plot with four blocks; the experimental factors
were irrigation (two levels, main plots) and nitro-
gen (two levels, subplots). With four blocks and
four treatments corresponding to each level of the
Cherfech Nabeul Nabeul
90–91 90–91 91–92
L1Y2 L3Y2 L3Y3
2 levels 2 levels 2 levels
2 levels 2 levels 2 levels
4 4
4 4
16 16 16
6 m×19 m 8 m×8 m 8 m×8 m
24 K. Latiri-Souki et al. / European Journal of Agronomy 9 (1998) 21–34
Table 2
Soil texture and soil chemical characteristics at the experimental locations
Texture
Clay Silt Sand
(0–2 mm) (2–50 mm) (50–2000 mm)
Cherfech 38 40 20
Nabeul 9 16 76
combination of the two experimental factors, the
plot was divided into 16 subplots of 6×19 m at
Cherfech and 8×8 m at Nabeul ( Table 1).
2.2.2. Irrigation
Shortly after sowing, an initial flood irrigation
(80 mm in year 1 at Cherfech, 45 and 38 mm at
Nabeul in year 2 and 3, respectively) was applied
to all the treatments in each experiment to
ensure uniform germination and emergence.
Subsequently, the water treatment was either no
irrigation (I−) or irrigation (I+), applied to return
the soil to field capacity and meet all the water
requirements of the crop. I+ plots were flood-
irrigated, water being applied whenever soil water
content was about two-thirds of the maximum
water storage capacity. At Cherfech, further irriga-
tion was applied only during the grain filling period
in both years with a single application of 100 mm
in Y1 and 165 mm in Y2. At Nabeul, a further
30–45 mm of irrigation was applied on each of six
occasions to the I+ treatment.
2.2.3. Nitrogen
Nitrogen treatment was either sufficient to give
a maximum yield (N+) based on previous experi-
ments or none (N−). In treatment N+, N was
applied by hand as ammonium nitrate at a concen-
tration of 33.33%. It was split into two equal
applications of 66 kg ha−1 when the crop had two
or three leaves and five or six leaves. In L1Y1, a
third amount was applied at anthesis
(66 kg ha−1) to replace N lost by leaching after
the high winter rainfall (December: 90 mm and
January: 150 mm).
pH Organic matter content C N
(%) (%) (g kg−1)
7.5–8 2.2 1.24 0.14
7.5–8 1.9 0.92 0.09
2.3. Environmental measurements
Climatic data were recorded at Cherfech and at
Nabeul using standard meteorological equipment
for rainfall, dry and wet air temperature and
sunshine hours.
2.3.1. Radiation and radiation use efficiency
Total radiation incident on the crop (R )
s
throughout the entire growing season was esti-
mated from bright sunshine hours (S) measured
on or close to the experimental sites, and from the
theoretical daylength (S ) and radiation at the top
0
of the atmosphere (R ) by using the Angström
a
formula (Doorenbos and Pruitt, 1984)
R =R [a+b.(S/S )],
s a 0
with coefficients provided by the National
Meteorological Office. For both sites, the coeffi-
cients are a=0.26, b=0.53 and a+b=0.79.
Intercepted radiation was measured at Nabeul with
a sunfleck ceptometer (Decagon, Pullman, WA)
using a 80-cm probe that was inserted just below
the height of the lowest leaves within the crop
canopy; three replicated measurement were made
in each subplot. Radiation use efficiency (RUE )
is calculated as either the above-ground dry matter
produced per unit of incident radiation or per unit
of intercepted radiation when measured.
2.3.2. Soil water content
Soil water content was measured either with the
gravimetric method for the upper 20 cm or with a
neutron probe to a depth of 120 cm. Access tubes
were placed in the upper third of each subplot.
Frequency of measurements was every 2 weeks and
 K. Latiri-Souki et al. / European Journal of Agronomy 9 (1998) 21–34
also on the day before irrigation and the day after
irrigation.
2.3.3. Crop water use
The measurement of crop water use was compli-
cated by heavy rainfall (in excess of 100 mm on a
maximum of two occasions in each year) and
drainage so that accurate measurements of evapo-
transpiration were difficult to obtain in the I+
treatment, especially at Cherfech, although the
estimate of ET in the I− plots and late in the
growing season was possible. The total water
supply was therefore equated with the amount of
water applied plus rainfall plus variation in soil
water content.
2.4. Crop measurement
Crops were sampled from December to May.
Growth was measured approximately every
2 weeks during vegetative growth and every week
during the grain filling period. Samples were taken
from 0.5 m of three adjacent rows (an area of
0.25 m2) and from 1 m of three adjacent rows
(0.5 m2) at anthesis and final harvest. The shoots
were cut off at the crown level and divided into
component parts for measurement. Leaf area was
measured with a planimeter (LI 3050-A, Li-COR
inc, Lincoln, NB, USA). At final harvest, grain
was separated from chaff, awns and straw and the
separate parts dried in an oven at 80°C for 48 h
before weighing.
Nitrogen content was measured at harvest on
Table 3
Crop management for the different location and years of experiment
L1Y1
Previous crop Tomato
Sowing 18-11-89 200 kg ha−1
450 grains m−2
Harvest Beginning of June
L3Y2
Previous crop Fallow
Sowing 11-12-90 220 kg ha−1
500 grains m−2
Harvest End of June
25
samples of grain, straw and chaff for each replicate
using the Kjeldahl method. The total amount of
N taken up by the crop was calculated by summing
the amount of N in each component.
2.5. Crop management
Plots were sown in mid-November to early
December with a seed density of 200 kg ha−1 at
Cherfech (which give 450 plants m−2) and
220 kg ha−1 at Nabeul (500 plants m−2): these
amounts were greater than usual in the area. The
plots were harvested in June. Agronomic treat-
ments were the same as those employed in the
region; they are summarised in Table 3.
2.6. Statistical analysis
Treatments were compared from the split-plot
designed experiment with four blocks, the main
plot being the I treatment and the subplots the N
treatments by analysis of variance obtained with
the statistical package Genstat (Genstat
Committee of the Statistics Department,
Rothamsted, 1992). Tests were conducted for signi-
ficance at 10, 5 and 1% probability.
3. Results
3.1. Weather conditions
The years of experiment were considered rainy,
although periods of drought occurred during the
L1Y2
Fallow
28-11-90 200 kg ha−1
450 grains m−2
End of June
L3Y3
Durum wheat
13-12-91 220 kg ha−1
500 grains m−2
End of June
26 K. Latiri-Souki et al. / European Journal of Agronomy 9 (1998) 21–34

growing season. Year 1 at Cherfech (L1) was
warmer than average, and the rainfall was slightly
below average (460 mm; average 478 mm). In year
2, rainfall was large during the period of vegetative
growth at L1, but grain filling had below average
rainfall at both Cherfech and Nabeul. In year 3,
the winter was drier at Nabeul than in year 2, but
the grain filling period was wetter.
3.2. Soil water and water use
The two soil types had very different water
storage capacities. At Cherfech, the maximum soil
water content was 414 mm to 1 m depth, whereas
at Nabeul, it was 220 mm to a depth of 1.20 m.
During drying at Cherfech, soil water content of
the profile did not decrease below 320 mm in the
I+ or I− treatments before anthesis. After anthe-
sis, the amount of water in treatment I− decreased
to about 270 mm, whereas the I+ treatment
remained at about 350 mm. At Nabeul, in the I−
treatment, the amount of water decreased pro-
gressively over the growing season from about
175 mm to 75 mm by final harvest. I+ was main-
tained between 150 and 200 mm throughout the
growth in both years (Fig. 1).
Water was extracted from the uppermost layers
of the profile first and from progressively deeper

Fig. 1. Water content change with calendar date for the different experimental treatments at Location 1 (a and b) and Location 3 (c
and d).
 K. Latiri-Souki et al. / European Journal of Agronomy 9 (1998) 21–34 27

soil as the profile dried at both locations (data not
given); it was extracted from deeper in the profile
in N+ treatments than in N− mainly at L3 and
at the end of the growing season. At Cherfech, the
soil water content did not differ significantly
between N treatments from sowing to harvest in
either year. At Nabeul, at the beginning of the
growing season, the soil water content in the profile
was slightly, but not significantly, larger in the
N+ treatments. In both years from March to
harvest, the soil water content in the profile was
decreased in the N+ treatments, resulting in less
water in the profile in the N+ treatments than in
the N− in the droughted treatment only. This
effect was significant only in year 3 and from the
end of March to harvest.
The total amount of water applied and used in
the different experiments varied between 280 mm
(L3Y3, I−N−) and 710 mm (L1Y2, I+N+)
( Table 4). Water use at L1 was very similar in I+
and I− treatments up to anthesis as the deficits
that developed were very small: during grain
growth, water use decreased as the deficit devel-
oped in the I− treatment compared to the I+
treatment with the droughted crop losing 12% and
16% less in year 1 and year 2, respectively, when
considering the whole growing season or 37 and
49% when considering the grain filling period only.
At Nabeul, the main variations were related to I
treatment. In I− treatments, the progressive
drying of the profile during vegetative as well as
grain growth decreased the amount of water used
by approximately 25% in Y2 and 36% in Y3.
3.3. LAI and PAR interception
3.3.1. LAI
The maximum values of LAI ranged from 1.3
(L3Y3, I−N−) to 7 (I+N+, L3Y2). The values
for the LAI in response to the N and I treatments,
averaged over location and year, are shown in
Fig. 2. The nitrogen application increased LAI
significantly during the whole growing season at
L3 and at L1Y1, but this effect was significant
only until stem elongation at L1Y2. At Nabeul, I
increased LAI and its duration significantly. At
the end of the season, the stems ears and awns
retained some chlorophyll, even when most of the
leaf area had yellowed. The length of the growing
season for the crops was judged according to the
retention of chlorophyll not according to leaf
area only.

Table 4
Water use for the different experimentsa

Experiment Treatment Water use for the different parts of the growing season

Sowing to anthesis (mm) Anthesis to harvest (mm) Sowing to harvest (mm)

L1Y1 I−N− 376 139 515

L1Y2 I−N− 464 130 594
L3Y2 I−N− 342 74 416
L3Y3 I−N− 202 78 280
L1Y1 I+N− 377 192 569
L1Y2 I+N− 481 218 699
L3Y2 I+N− 341 218 559
L3Y3 I+N− 298 150 448
L1Y1 I−N+ 387 116 503
L1Y2 I−N+ 485 102 587
L3Y2 I−N+ 354 70 424
L3Y3 I−N+ 218 73 291
L1Y1 I+N+ 375 215 590
L1Y2 I+N+ 476 234 710
L3Y2 I+N+ 341 218 559
L3Y3 I+N+ 304 145 448

aWater use: rain, irrigation and variation in soil water content are considered, but drainage is neglected.
28 K. Latiri-Souki et al. / European Journal of Agronomy 9 (1998) 21–34

Fig. 2. Leaf Area Index with calendar date for the different experimental treatments at Location 1 (a and b) and Location 3 (c and d ).

3.3.2. Incident PAR radiation ranged from 978 MJ m−2 (L1Y1)

The incident PAR received by the crop is calcu- to 1152 MJ m−2 (L3Y3). At Nabeul, drought
lated for the number of days for which the decreased the growing season by reducing the grain
crop was green. The total accumulated incident filling period, and therefore, the amount of PAR

Table 5
Incident PAR and incident radiation use efficiency for the different treatments and the different years of experiment

Sowing to anthesis Incident PAR (MJ−1) Incident radiation use efficiencies (g MJ−1)

Anthesis to harvest I−N− I+N+ I+N− I+N+

I− I+

L1Y1 527 452 452 1.54 1.82 1.49 1.83

L1Y2 670 661 661 0.80 0.93 0.77 0.86
L3Y2 603 442 511 0.65 1.02 1.36 1.83
L3Y3 666 385 486 0.60 1.19 0.98 1.32
 K. Latiri-Souki et al. / European Journal of Agronomy 9 (1998) 21–34 29

Table 6
Intercepted PAR, intercepted PAR use efficiency (measured at L3) for the different treatments and averaged over the different years
of experiment

Intercepted Percentage of incident PAR Efficiency of

PAR (MJ ) intercepted by the crop 1)
intercepted PAR (g MJ−

L3Y2 I−N− 560 54 1.2

L3Y2 I−N+ 771 74 1.4
L3Y2 I+N− 808 73 1.9
L3Y2 I+N+ 930 83 2.2
L3Y3 I−N− 410 39 1.5
L3Y3 I−N+ 592 56 2.1
L3Y3 I+N− 668 58 1.7
L3Y3 I+N+ 751 65 2.0

incident on the crop during the grain filling period
was smaller by 17%. In the irrigated crop, the lack
of nitrogen averaged over all factors did not affect
the total PAR compared to the N+ treatments
( Table 5).
nificantly increased dry matter production at both
locations by increasing both straw and grain dry
matter. However, the proportional increase in
straw dry matter was slightly larger, resulting in a
small reduction of harvest index with high N

3.3.3. Intercepted radiation

The amount of PAR intercepted by the crop
was measured only at Nabeul. The greater LAI
with applied I and N increased PAR interception,
which varied from between 410 MJ m−2 (L3Y3,
I−N−), corresponding to 39% of incident radia-
tion measured over the crop life, and
930 MJ m−2 (L3Y2, I+N+), corresponding to
83% of incident radiation ( Table 6).

3.4. Dry matter production, grain yield and

environment
(a)
The total above-ground biomass production was
greatest in the I+N+ treatments averaged over
all years and locations [Fig. 3(a)]. Treatments that
received no addition of N fertiliser or irrigation,
after that given to obtain good germination
(I−N−), produced the smallest total dry matter
and grain yields. This was particularly so in the
sandy, low fertility soil at Nabeul.
Total dry matter increased with the total amount
of water (irrigation+rainfall+variation in soil
water content) available to the crop [Fig. 3(a)].
Grain yield [Fig. 3(b)] responded in the same way, (b)
so the harvest index was almost constant. It was Fig. 3. (a) Total dry matter against water supply for the two
44.4% in the I− and I+ treatments averaged over levels of nitrogen. (b) Grain yield against water supply for the
all other conditions. Application of nitrogen sig- two levels of nitrogen.
30 K. Latiri-Souki et al. / European Journal of Agronomy 9 (1998) 21–34

(43.2% compared to 45.5% for N−, difference not

statistically significant). For each amount of water
applied to the crop, more dry matter was produced
in the treatment with N than that without N
[Fig. 3(a)]. Both the smallest and largest values of
biomass production were observed at Nabeul (L3,
sandy soil ), whereas at Cherfech (L1, clay soil ),
the differences were smaller, and only an increase
related to N treatment was observed: the increase
related to I was small and not significant.
(a)
3.5. Radiation and water use efficiencies

The incident radiation use efficiency (Table 5)

ranged from 0.6 g MJ−1 (I−N− at L3Y3) to
1.83 g MJ−1 (I+N+ at L1Y1 and L3Y2). For
the whole growing season, the intercepted RUE
efficiency was between 1.2 (I−N−, L3Y2) and
2.2 g MJ−1 (I+N+, L3Y2). Treatments deficient
in water and applied nitrogen had the lowest
efficiencies, whereas treatments with applied I and
N had the largest: Irrigated and fertilised crops
provide an estimate of the maximum potential
efficiencies: 1.83 g MJ−1 for Karim at Nabeul and
at Cherfech.
Intercepted RUE increased linearly with N
uptake [Fig. 4(a)]. RUE and normalized water use
efficiency (dry matter per unit of water per unit of
vapour pressure deficit) are related [Fig. 4(b)],
reaching a plateau at large values of water use
efficiency.
3.5.1. Water use efficiency
Water use efficiency is considered for different
periods of time, sowing to anthesis, anthesis to
harvest and sowing to harvest. From sowing to
anthesis, nitrogen always increased water use effi-
ciencies when considering either total water
received, water evaporated or water evaporated
per unit of vapour pressure deficit (the normalised
water use efficiency). Also, efficiencies were larger
without irrigation than with irrigation when con-
sidering the same amount of nitrogen applied
[Fig. 5(a)]. During the grain filling period, effi-
ciencies decreased with increasing water or even
had negative values, e.g. for I−N+ at L3Y2 as a
slight decrease in total dry matter occurred
between anthesis and harvest. For the other treat-
(b)
Fig. 4. (a) Intercepted radiation use efficiency (RUE ) averaged
over irrigation against N uptake. (b) Logarithmic regression of
Intercepted Radiation Use Efficiency against normalized water
use efficiency.
ments, years and locations during the grain filling
period, efficiencies were increased by applied nitro-
gen [Fig. 5(b)]. At L3, total dry matter per unit
of water evaporated per unit of vapour pressure
deficit ranged from 0.007 mb to 0.018 mb
[Fig. 5(c)]. It was increased by nitrogen applica-
tion, and this effect was more important in year 3
(I−N−: 0.009 mb, I−N+: 0.018 mb) than in
year 2 (I−N−: 0.007 mb, I−N+: 0.011 mb).
Also, during the grain filling period at L1, WUE
was larger with applied nitrogen than without: the
greatest value was at L1Y1 for treatment I−N+
(0.039 mb).
3.6. Nitrogen content and use efficiency
For the whole crop, a lack of N in the I−
treatment substantially reduced the percentage N
in the whole crop throughout growth, whereas
 K. Latiri-Souki et al. / European Journal of Agronomy 9 (1998) 21–34 31

Nitrogen use efficiency (NUE: grams of dry

matter per gram of N absorbed ) ranged from 58
(L3Y3, I+N−) to 102 (I−N−, L3Y3) ( Table 7).
Averaged over the locations and years, it was
smallest in the crops with the greatest dry matter
and greatest for the crops receiving no water
or nitrogen application. Irrigation particularly
decreased the NUE overall by 13%, and N applica-
tion decreased it by 6%.

(a)
4. Discussion

Our results have shown that on both soils,

(b)
(c)
Fig. 5. (a) Normalized water use efficiency against water supply
with (N+) or without (N−) applied nitrogen for the period
from sowing to anthesis. (b) Normalized water use efficiency
against water supply with (N+) or without (N−) applied nitro-
gen for the period from anthesis to harvest. (c) Normalized
water use efficiency against water supply with (N+) or without
(N−) applied nitrogen for the whole growing season.
application of N stimulated the growth of LAI
and green stem and ear areas and increased crop
production. Despite the fact that LAI decreased
with deficient N, the green ear area was maintained
for the same time as that in the N+ treatments
so that the total period of crop growth was consid-
ered the same with and without N: hence, the
incident radiation was the same for crops with
both N treatments. This is different from other
studies (Gajri et al., 1993) where N application
gave not only a larger LAI and ear area but also
maintained the duration of green area for longer
than N-deficit crop, thus affecting the total energy
available to the crop.
Irrigation increased the LAI and green ear area
and their duration compared to droughted treat-
ments in both years and at both sites. Hence, there
was no interaction between I and N on the period
for which the crop remained green. Other studies
(Gajri et al., 1993) have shown interactions with
application of N increasing the duration of the
leaf area and green crop under irrigation compared
to deficient N. Droughted crops generally mature
earlier, depending on the severity of the stress,
Table 7
1 ) for the different treatments
Nitrogen use efficiency (g g−
I−N− I−N+ I+N− I+N+

L1Y1 84 82 79 68
with irrigation, the crop receiving no applied N L1Y2 83 80 90 80
had only a slightly smaller percentage N than the L3Y2 85 83 69 79
L3Y3 102 74 59 63
crop with applied N, but not to the same extent
Average 88.5 79.8 74.3 72.5
as the I−N− treatment.
32 K. Latiri-Souki et al. / European Journal of Agronomy 9 (1998) 21–34
temperature, etc. Under very dry conditions, there
is often a considerable advance in senescence,
particularly with abundant N compared to defi-
cient (Anderson, 1985; Papastylianou, 1995).
Therefore, there is an interaction between N and
I. This is the basis for the generally held belief
that application of N has negative effects on pro-
duction under dry conditions.
The total amount of incident radiation judged
on the basis of the duration of green area (from
leaves, ears, awns or stems) is an inexact way of
judging how effective the crop is in using any
available energy so the amount of crop interception
of energy is important.
The proportion of incident energy absorbed by
the crop at Nabeul shows the impact of applying
N and I. Applied N increased interception by 13%,
and irrigation by 14%, averaged over 2 years and
other factors. Thus, combining I and N increased
the interception of energy substantially due to the
larger LAI, but as the crop matured and leaf area
senesced, a smaller proportion of the incident
energy was intercepted, and a greater proportion
of the intercepted energy was captured by ears and
awns: these are less efficient in converting energy
to dry matter.
Increased interception of radiation is the main
driving force for the improvement in crop biomass
production. This explains, in part, how crop
growth was affected by the availability of N and
I. The effects were large: maximum above ground
dry matter production of 20 t ha−1 at Nabeul and
18 t ha−1 at Cherfech was large for a durum wheat
crop grown for 7 months in the relative warm, dry
conditions of Tunisia. In the very cool moist
environment of Britain, winter wheat crops grown
for 10 months may produce 25 t ha−1 of above-
ground dry matter (Russel and Wilson, 1994).
Such large biomass production in Tunisia reflects
the abundant radiation, even in the winter period,
so the application of I and N with good crop
management enables large rates of production and
gives substantial biomass at final harvest. Under
stress conditions, with no I or N applied, with the
crop using only stored water and rainfall and N
available in the soil, production depended on the
soil type. At Cherfech, the high fertility and water
holding capacity of the soil enabled the crops not
supplied with I and N to produce 80% of the dry
matter of the control. At Nabeul, the effect of
deficiency of I and N (I−N−) was to decrease
crop production by 65% compared to the control
(I+N+).
The reason for the beneficial effects of I and N
relate to the development of the crop and especially
to LAI and the impact this has on radiation
interception, as discussed earlier. However, the
consequences of I and N supply on the efficiency
with which radiation is used by the crop are also
of importance ( Table 6 and Fig. 4). The efficiency
of radiation conversion to dry matter, the RUE,
was substantially increased (45%) by applied nitro-
gen fertilizer. The percentage N in crop dry matter
during the rapid period of vegetative growth
(approximately maximum LAI ) was about 3% in
the N+ treatments irrespectively of I, whereas it
is about 2% without N, so an approximately 50%
increase of N% in dry matter results in a larger
increase in RUE. Thus, N increased radiation
interception by 24% (i.e. total energy available for
growth) and radiation use efficiency by 22%
(Table 6). Under the conditions of these experi-
ments in different years and at two locations, it is
clear that N application has a substantial positive
impact on durum wheat production, when
averaged over other conditions. This is shown in
Fig. 3 where the total dry matter at any water
application is increased; the effect is smaller with
ample water than in droughted conditions, where
applied N can double the total dry matter. It is
important that the N is applied early in the crop
growth so that leaf development and organ differ-
entiation can proceed without limitation (Lawlor
et al., 1981; Latiri-Souki et al., 1992; Giunta et al.,
1993). Such a substantial effect on total dry matter
in the experiments under the range of conditions
experienced suggests that the advantage of N appli-
cation will be seen in practical agriculture in semi-
arid conditions.
When water is limiting crop production, conser-
vation and efficient use of water are important
considerations for agriculture, in addition to the
need for production. Although the application of
N increased the LAI in our experiments, it did not
significantly increase the water loss from the crops.
In the early part of the growing season, the
 K. Latiri-Souki et al. / European Journal of Agronomy 9 (1998) 21–34
increased LAI with N+ protected the soil surface
from evaporation and resulted in a slightly greater
amount of soil water in the root zone than with
N−; during the later part of the growing season,
the N+ crops evaporated this retained water and
some additional water so that the total water
retained in the soil was slightly less than with N−
treatment ( Fig. 1). Hence, application of N to
crops had only a small effect on crop water use,
and the benefits of N application clearly out-
weighed any disadvantages.
As a consequence of the increased dry matter
production with irrigation but the larger relative
water use, the WUE of the irrigated crops was
smaller than the WUE of the droughted crops.
Without applied N, the difference was only slight.
However, in very dry conditions at Nabeul, appli-
cation of N greatly increased WUE, in fact more
than doubled it, as the amount of applied water
decreased. During the grain filling period, the
WUE decreased with greater water supply, and
there was no difference between N treatments. This
period is characterised by decreasing green leaf
area and greater contribution of ears, awns and
stems to radiation interception. Assimilate pro-
duction decreases, and there is a remobilisation of
assimilates from vegetative organs to grains
(Gallagher et al., 1975). Thus, the balance of dry
matter growth and water use seen in the vegetative
period is greatly altered during grain filling. Hence,
WUE during the grain filling period is not compa-
rable to that in vegetative growth.
Irrigation and nitrogen applications to crops
growing under conditions where these resources
are inadequate to meet the genetic potential for
growth are well known to improve plant and yield
production (Smith and Harris, 1981). Our study
shows that for durum wheat, production may be
doubled in dry conditions with poor water supply
by applying N fertilizer, providing that other nutri-
ents are adequate to meet the needs of the more
rapidly growing crop and that crop management
(pest and disease control, etc.) is commensurate.
The effect of N from our studies is applicable to
very different soil types, although the extent of the
response may differ depending on soil organic
matter and water holding capacity. In very dry
years, water is the limiting factor for plant growth,
33
and under these conditions, root growth is
restricted, and fertilizer nitrate is not effectively
dissolved in the soil water so applied N will not
be absorbed efficiently. In extreme droughts on
very dry soils, water may be expected to dominate
crop growth: any benefit of applied N would not
be seen. The frequency of such droughts is approxi-
mately 20% in the semi-arid areas of Tunisia on
which durum wheat is grown ( El Amami, 1984).
Therefore, the advantage of N application would
be smaller on overall production from those area,
but still the long term net advantage would be
considerable.
Acknowledgment
Partial funding from the British Council and
from the International Foundation for Science is
gratefully acknowledged. IACR receives grant-
aided support from the Biotechnology and
Biological Sciences Research Council of the United
Kingdom.
References
Anderson, W.K., 1985. Grain yield response of barley and
durum wheat to split nitrogen applications under rainfed con-
ditions in a Mediterranean environment. Field Crops Res.
12, 191–202.
Aubry, C., Latiri-Souki, K., Dore, T., Griner, C., 1994.
Diagnostic des facteurs limitants du rendement du blé dur en
parcelles d’agriculteurs dans une petite région semi-aride en
Tunisie. Agronomie 14, 213–227.
Brown, S.C., Keatinge, J.D.H., Gregory, P.J., Cooper, P.J.M.,
1987. Effects of fertilizer, variety and location on barley pro-
duction under rainfed conditions in Northern Syria. I. Root
and shoot growth. Field Crops Res. 16, 53–66.
Chaabouni, Z., 1983. Caractéristiques physiques des sols de la
parcelle expérimentale de Oued Souhil, Nabeul. Centre de
Recherche du Génie Rural, Tunis.
Cooper, P.J.M., Gregory, P.J., Tully, D., Harris, H.C., 1987.
Improving water use efficiency of annual crops in the rainfed
farming systems of West Asia and North Africa. Exp. Agric.
Farming Syst. Ser.-5 23, 113–158.
Dann, P.R., 1969. Response of wheat to phosphorus and nitro-
gen with particular reference to ‘haying-off ’. Austral. J. Exp.
Agric. Anim. Husb. 9, 625–629.
Day, W., Lawlor, D.W., Legg, B.J., 1981. The effects of drought
on barley: soil and plant water relations. J. Agric. Sci. Camb.
96, 61–77.
34 K. Latiri-Souki et al. / European Journal of Agronomy 9 (1998) 21–34
Doorenbos, J., Pruitt, W.O., 1984. Crop Water Requirements.
FAO Irrigation and Drainage, 24.
El Amami, S., 1984. Sécheresse et planification de la production
céréalière en Tunisie. Centre de Recherche du Génie Rural,
Tunis.
Evans, L.T., 1993. Crop Evolution, Adaptation and Yield.
Cambridge University Press, Cambridge.
Fisher, R.A., 1981. Optimizing the use of water and nitrogen
through breeding of crops. Plant and Soil 58, 249–278.
Gajri, P.R., Prihar, S.S., Arora, V.K., 1993. Interdependence
of nitrogen and irrigation effects on growth and input-use
efficiencies in wheat. Field Crops Res. 31, 71–86.
Gallagher, J.N., Biscoe, P.V., Hunter, B., 1975. Barley and its
environment. V. Stability of grain weight. J. Appl. Ecol. 12,
319–336.
Genstat 5 Committee Of The Statistics Department, 1992.
Genstat 5 Reference Manual. Rothamsted Experimental
Station. Oxford University Press, Oxford.
Giunta, F., Motzo, R., Deidda, M., 1993. Effect of drought on
yield and yield components of durum wheat and triticale in
a Mediterranean environment. Field Crops Res. 33, 399–409.
Gonzales Ponce, R., Salas, M.L., Mason, S.C., 1993. Nitrogen
use efficiency by winter barley under different climatic condi-
tions. J. Plant Nutr. 16, 1249–1261.
Greenwood, D.J., 1982. Modelling of crop response to nitrogen
fertilizer. Phil. Trans. R. Soc. B 296, 351–362.
Greenwood, D.J., Gastal, F., Lemaire, G., Draycott, A.,
Millard, P., Neeteson, J.J., 1991. Growth rate and % N of
field grown crops: Theory and experiments. Ann. Botany
67, 181–190.
Grindlay, D.J.C., 1997. Towards an explanation of crop nitro-
gen demand based on the optimization of leaf nitrogen per
unit leaf area. J. Agric. Sci. Camb. 128, 377–396.
Kassam, A.H., 1981. Climate, soil and land resources in North
Africa and West Asia. Plant and Soil 58, 1–29.
Latiri-Souki, K., Aubry, C., Dore, T., Sebillote, M., 1992.
Elaboration du rendement du blé dur en Tunisie I—Relations
entre composantes du rendement sous différentes régimes de
nutrition azotée et hydrique. Agronomie 12, 31–43.
Lawlor, D.W., 1995. Photosynthesis, productivity and environ-
ment. J. Exp. Botany 46, 1449–1461.
Lawlor, D.W., Day, W., Johnston, A.E., Legg, B.J., Parkinson,
K.J., 1981. Growth of spring barley under drought: crop
development, dry matter accumulation and nutrient content.
J. Agric. Sci. 96, 167–186.
Lawlor, D.W., Boyle, F.A., Keys, A.J., Kendall, A.C., Young,
A.T., 1988. Nitrate nutrition and temperature effects on
wheat: a synthesis of plant growth and nitrogen uptake in
relation to metabolic and physiological processes. J. Exp.
Botany 39, 329–343.
Legg, B.J., Day, W., Lawlor, D.W., Parkinson, K.J., 1979. The
effects of drought on barley growth: models and measure-
ments showing the relative importance of leaf area and photo-
synthetic rate. J. Agric. Sci. 92, 703–716.
Monteith, J.L., 1977. Climate and the efficiency of crop pro-
duction in Britain. Phil. Trans. R. Soc. B 281, 277–294.
Monteith, J.L., Elston, J., 1993. Climatic constraints on crop
production. In: Fowden, L., Mansfield, T., Stoddart, J.
( Eds.), Plant Adaptation to Environmental Stress. Chapman
& Hall, London, pp. 3–18.
Papastylianou, I., 1995. Yield components in relation to grain
yield losses of barley fertilized with nitrogen. Eur. J. Agron.
4, 55–63.
Papy, F., 1979. Analyse du comportement des cultures de blé
dur et d’orge dans différentes régions céréalières du Maroc à
travers leur réaction aux variations interannuelles des régimes
pluviométriques. CR Séances Acad. Agric. France 65,
231–247.
Ritchie, J.T., 1983. Efficient water use in crop production:
Discussion on the generality of relations between biomass
production and evapotranspiration. In: Taylor, H.M.,
Jordan, W.R., Sinclair, T.R. ( Eds.), Limitations to Efficient
Water Use in Crop Production. American Society of
Agronomy, Madison, WI, pp. 29–43.
Russel, G., Wilson, G.W., 1994. An agro-pedo-climatological
knowledge base of wheat in Europe. JAC 158.
Shepherd, K.D., Cooper, P.J.M., Allan, A.Y., Drennan,
D.S.H., 1986. Growth, water use and yield of barley in
Mediterranean-type environments. J. Agric. Sci. Camb.
108, 365–378.
Sinclair, T.R., Horie, T., 1989. Leaf nitrogen, photosynthesis,
and crop radiation use efficiency: a review. Crop Sci. 29,
90–98.
Smith, R.C.G., Harris, H.C., 1981. Environmental resources
and restraints to agricultural production in a Mediterranean
type environment. Plant and Soil 58, 31–57.
Tanner, C.B., Sinclair, T.R., 1983. Efficient water use in crop
production: Research or re-search? In: Taylor, H.M., Jordan,
W.R., Sinclair, T.R. ( Eds.), Limitations to Efficient Water
Use in Crop Production. American Society of Agronomy,
Madison, WI, pp. 29–43.
Tardieu, F., Katerji, N., Bethenod, O., Zhabg, J., Davies, W.J.,
1991. Maı̈ze stomatal conductance in the field: its relationship
with soil and plant water potentials, mechanical constraints
and ABA concentration in the xylem soap. Plant Cell
Environ. 14, 121–126.
Thorne, G.N., Welbank, P.J., Widdowson, F.V., Penny, A.,
Todd, A.D., Weit, A.H., 1988. Contrast between sandy and
clay soils in the effects of various factors on the growth,
nitrogen uptake and yield of winter wheat in three years.
J. Agric. Sci. 110, 119–140.
UNDP-UNESCO, 1970. Research and training on irrigation
with saline water (1962–1969) Technical report. United
Nations Development Programme, United Nations
Educational Scientific and Cultural Organisation, Paris.
van Duivenbooden, N., de Wit, C.T., van Keulen, H., 1996.
Nitrogen, phophorus and potassium relations in five major
cereals reviewed in respect to fertilizer recommendations
using simulation modelling. Fertilizer Res. 44, 37–49.
van Keulen, H., Seligman, N.G., 1987. Simulation of water use,
nitrogen nutrition and growth of a spring wheat crop. Pudoc,
Wageningen.