1 s2.0 S092666901530234X Main

Industrial Crops and Products 76 (2015) 494508
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Industrial Crops and Products

journal homepage: www.elsevier.com/locate/indcrop
Osmoregulation-mediated differential responses of eld-grown

fennel genotypes to drought
Ehsan Askari , Parviz Ehsanzadeh
Department of Agronomy and Plant Breeding, College of Agriculture, Isfahan University of Technology, Isfahan, 84156-83111, Iran
a r t i c l e
i n f o
Article history:
Received 11 January 2015
Received in revised form 8 May 2015
Accepted 9 July 2015
Available online 3 August 2015
Keywords:
Foeniculum vulgare
Grain yield
Essential oil
Irrigation
Osmolytes
a b s t r a c t
This study was aimed at examination of physiological and agronomic responses of fennel (Foeniculum
vulgare Mill.) to water decit. Twelve fennel genotypes namely Kashan, Urmia, Hamadan, Kerman,
Shiraz, Birjand, Mashhad, Ardabil, Bushehr, Avicenna, Isfahan, and Yazd were subjected to four
levels of irrigation (irrigation after 35%, 55%, 75%, and 85% depletion of available soil water) in a eld study
in two years. Leaf water potential, relative water content (RWC), proline, total soluble sugars, chlorophyll
a (Chl a), b (Chl b), total (Chl a + b), a/b (Chl a/b) along with dry mass (DM), seed yield and its attributes and
seed essential oil content were measured. Water deprivation left signicant effects on all characteristics,
i.e. in contrast to seed essential oil content and harvest index, the rest of the attributes were decreased
signicantly with drought intensication. Varietal differences in response to drought were meaningful
for most of the traits; i.e. genotypic variations for DM, grain yield and stress susceptibility index (SSI)
were consistent with differences among the genotypes in physiological traits such as leaf water potential,
RWC, proline, soluble sugars and chl content. Genotypes were discriminated according to their response
to drought and SSI. Drought tolerant genotypes (Yazd, Kerman, Mashhad and Shiraz) exhibited a
greater capacity for accumulation of osmotic solutes associated with higher leaf water potential and
RWC, compared to drought sensitive genotypes (Ardabil, Avicenna, Hamadan and Birjand). From
our ndings, fennel could be appreciated as a promising species in potentiating alternative industrialmedicinal crops in the face of the eminent challenge of water scarcity in arid and semi-arid climatic
regions.
2015 Elsevier B.V. All rights reserved.
1. Introduction
Essential oils from different plants are valuable complex products that various pharmaceutical and biological activities from
antibacterial to antifungal, antiviral and anticancer are attributed
to them (Raut and Karuppayil, 2014). Fennel is a perennial essential
oil-bearing plant with traditional uses in medicine and human consumption that is grown in arid and semi-arid regions of the world
(Ashraf and Akhtar, 2004), including Iran. Fennel seeds (fruits) and
their essential oil are used in pharmaceutical and cosmetic industries and food production throughout the world (Ehsanipour et al.,
2012). Fennels products have proven useful in the treatment of a
variety of complaints including diabetes, chronic coughs and kidney
stones (Barros et al., 2009). Several parts of fennel including roots,
pseudo bulbs, young leaves, shoots and seeds contain essential oil
(Sarkheil et al., 2008).
Corresponding author. Fax +98 311 391 3453.

E-mail address: ehsanaskariii@gmail.com (E. Askari).
http://dx.doi.org/10.1016/j.indcrop.2015.07.010
0926-6690/ 2015 Elsevier B.V. All rights reserved.
Water remains the single most important factor affecting world

food security, e.g. it limits crop production particularly in dry
regions of the world, where they are characterized by regular and
prolonged drought occurrences (Fageria et al., 2006). Therefore, the
foremost abiotic stress is water decit in many parts of the world,
including countries of Middle East. In fact, about 28% of the world
land area is estimated to face with limited crop productivity due
to varying degrees of drought stress. Water decit affects a vast
range of plant characteristics including plant morphology, physiology and biochemistry (Mohamed and Abdu, 2004) and, therefore,
productivity. There is a dire need to improve the production of
water-efcient plants in arid and semi-arid parts of the world to
tackle the progressive water scarcity. Considering the lower water
requirement at least in some medicinal plants, compared to many
non-medicinal crop plants, increasing the areas under cultivation
of these plants seems advantageous for arid and semi-arid parts of
the world.
Vegetative and reproductive growth and physiological process
involved in, herb and seed yield, quantity and quality of essential
oil of fennel and other medicinal plants depend to a great extent
E. Askari, P. Ehsanzadeh / Industrial Crops and Products 76 (2015) 494508
495
Table 1
A synopsis of physical and chemical properties of the experimental soil.
Bulk density
(g cm3 )
pH
Water-holding
capacity at eld
capacity (g kg1 )
Organic C content (g kg1 )
Total N (mg kg1 )
Available P (mg kg1 )
Available K (mg kg1 )
EC (dS m1 )
1.39
7.4
230
3.6
450
25.5
225
1.6
on factors such as climate, environmental stresses, cultivar, sowing date, harvest date, diseases, weed pressure and management
practices (Bowes and Zheljazkov, 2005). Since fennel can be grown
in marginal land and is considered tolerant to various stresses,
it might be a suitable medicinal crop for drought-prone environments. But scientic data on the response of this medicinal plant to
watering regimes is scarce. In order to address this gap in knowledge, the present study was carried out to evaluate the efcacy
of four irrigation regimes on essential oil, some physiological and
growth characteristics, yield and yield components of twelve fennel
genotypes.
2. Materials and methods
2.1. Experimental setup, plant material and growth conditions
A two-year (i.e. 2011 and 2012) eld experiment was carried
out at the Lavark Research Farm of Isfahan University of Technology, located in Najaf Abad (32 32 N, 51 23 E, 1630 m above mean
sea level, mean annual temperature 14.5 C, and 140 mm mean
annual precipitation), Iran. The daily minimum (T-min) and maximum (T-max) air temperature, minimum (RH-min) and maximum
(RH-max) relative humidity and reference evapotranspiration (ET0 )
during the growing season in 2011 and 2012 are shown in Fig. 1AC,
respectively.
The objective of the study was to investigate the effects of four
irrigation regimes (consisting irrigation after 35%, 55%, 75% and
85% depletion of available soil water) on 12 fennel genotypes, collected from different regions in Iran, namely Urmia, Hamadan,
Kerman, Shiraz, Birjand Yazd Ardabil, Avicenna, Kashan,
Mashhad, Bushehr and Isfahan. The experiment was designed
as a three replicate split-plot randomized complete block, in which
main plots consisted of the four irrigation regimes and subplots
consisted of the 12 fennel genotypes.
The seeds were treated with 5% sodium hypochlorite solution for
1 min, then repeatedly rinsed with distilled water to remove excess
sodium hypochlorite solution. In order to obtain adequate and uniform seedlings in the eld, healthy seeds were pre-germinated in
54 28 cm propagation trays containing coco peat and grown in
a green house. Three-weeks-old seedlings (i.e. 4-leaf stage) were
transplanted into the eld plots in late March 2011. Each experimental unit (subplot) consisted of ve 2-m long rows with 0.5 m
spacing between rows and 0.2 m spacing between plants in the
same row. Therefore, the seedlings were sown at an approximate
planting rate of 10 plants m2 in each plot.
The soil characteristics (Fine Loam Typical Haplargid) are given
in Table 1. Since the soil was decient in nitrogen, a urea fertilizer
containing 46% of N was given uniformly at a 150 kg ha1 basis to
the soil prior to sowing.
2.2. Irrigation regimes
The transplanted plants in all experimental plots were irrigated
uniformly when 35% of available soil water (ASW) was depleted for
ve weeks, then when the plants were approximately 20 cm tall
watering regimes were applied and continued to approximately
70% physiological maturity, i.e. late September 2011. For the 2012
year the plots were given two uniform irrigations on March and the
Table 2
The root zone depth for each genotype (measured three times in each season in May,
July and September) during the growing seasons of 2011 and 2012.
Genotype
Urmia
Hamadan
Kerman
Shiraz
Birjand
Yazd
Ardabil
Avicenna
Kashan
Mashhad
Bushehr
Isfahan
Mean
Root zone depth (cm) 2011
Root zone depth (cm)
2012
May
July
September
May
July
September
24
21
20
30
25
21
20
37
30
34
36
31
27.4
44
34
33
48
45
39
34
52
49
50
54
47
44.1
52
44
42
59
53
45
43
67
61
66
67
59
54.9
56
49
46
71
59
49
46
82
71
79
80
74
63.5
68
59
56
86
71
59
56
94
86
92
95
87
75.7
79
68
64
94
84
68
68
104
98
101
105
99
86.0
irrigation regimes were applied on early April and continued until

mid-September 2012.
Total ASW is dened as amount of the soil water in the root zone
between eld capacity and the permanent wilting point. ASW was
calculated based on Eq. (1).
ASW = (WFC WWP ) Bd V
(1)
where WFC is the gravimetric soilwater content (%) at eld capacity, WWP the gravimetric soilwater content (%) at the permanent
wilting point, Bd the bulk density of the soil (g cm3 ) and V is the
volume of soil layer in the root zone (m3 ). Root zone depth for all
genotypes was measured three times during each growing season.
At each time point, the measured root zone depths were averaged over the genotypes and the watering treatments were applied
according to the calculated mean root zone depth (Table 2). The
fraction of ASW that a crop can readily extract from the root zone
without suffering water stress is dened as the readily available soil
water (RAW) and was calculated according to Eq. (2) (Allen et al.,
1998).
RAW = ASW
(2)
The factor varies for different plants from 0.3 for shallowrooted crops at high rates of plant evapotranspiration, ETc (>8 mm
day1 ) to 0.7 for deep rooted crops at low rates of ETc (<3 mm
day1 ) (Allen et al., 1998). The factor was used to estimate the
required time of irrigation to prevent water stress. At low rates of
ETc (5 mm day1 ), the value of is recommended to be 0.4 for
parsnip, another member of Apiaceae family. Thus, the latter value
was used for fennel in this study. The fraction is dened as a
function of the evaporation power of the atmosphere based on Eq.
(3).
= rec + 0.04(5 ETc )
(3)
where rec is the recommended value for parsnip, and is the

adjusted value for evaporation demand of the atmosphere. Following determination of , irrigation treatments were scheduled based
on the maximum allowable depletion (MAD) percentage of ASW
(Kramer and Boyer, 1995). Irrigation was applied when 35%, 55%,
75% and 85% of the ASW was depleted from the root zone. The latter levels of irrigation were, arbitrarily, designated as the control,
496
2011
2012
T-max
(A)
45
40
40
Temperature ( C)
T-min
45
35
35
30
30
25
25
20
20
15
15
10
10
RH (%)
(B)
ET0 (mm)
(C)
RH-min
T-min
RH-max
T-max
RH-min
100
100
80
80
60
60
40
40
20
20
15
15
13
13
11
11
-1
-1
RH-max
I1 (35% Depletion of ASW)
(D)
1200
Cumulative applied water (mm)
Cumulative applied water (mm)
1000
900
800
700
600
500
400
300
200
100
0
Apr
May
Jun
Jul
Aug
Sep
1000
800
600
400
200
0
Apr
May
Jun
Jul
Aug
Sep
Fig. 1. The daily minimum (T-min) and maximum (T-max) air temperature (A); minimum (RH-min) and maximum (RH-max) relative humidity (B); reference evapotranspiration (ET0 ) (C) and cumulative applied water for irrigation regimes (I1, I2, I3 and I4 = 35, 55, 75 and 85% depletion of available soil water, respectively) (D), during the growing
seasons in 2011 and 2012.
mild, moderate and severe levels of drought stress, respectively. A

TDR probe (TDR Trase System, Model 6050XI; Soil Moisture, Santa
Barbara, CA) was used to measure soil water content two days after
irrigation and continued up to one day prior to the next irrigation.
The depletion of the available soil water based on the soil water
potential was determined by a soil moisture release curve. The
volume of irrigation water (Virrig ) necessary to increase the water
content in the root zone depth to eld capacity was calculated using
Eq. (4).
Virrig
ASW f
=
Ea
(4)
where f is the fraction of ASW (35%, 55%, 75% and 85%) that can
be depleted from the root zone, and Ea is the irrigation efciency
(%). Irrigation efciency was assumed to be 70% throughout the

growing season (Tafteh and Sepaskhah, 2012). The irrigation water
was applied with a pipe and the volume was measured with a ow
meter. The cumulative applied water (mm) for irrigation regimes
(I1 , I2 , I3 and I4 = 35, 55, 75 and 85% depletion of available soil
water, respectively) during the growing seasons in 2011 and 2012
is shown in Fig. 1D.
2.3. Measurement of leaf water relations, osmolytes and
photosynthetic pigments
At 50% owering stage, leaf water potential and relative water
content, and leaf proline, soluble sugars and chlorophyll contents
of three plants per experimental unit were measured.
The mid-day leaf water potential was determined using a

portable pressure chamber instrument (PMS Model 600, USA). On
a sunny day, second fully developed upper leaves were excised at
the petioles close to leaf collars. The chamber was pressurized with
compressed air until the tissue water was returned to the open end
of the petiole and could be seen in the cut surface. Then the measured balance pressure was explained as the leaf water potential
and expressed as Mega Pascals (MPa). A mean of three measurements were reported for each plot.
RWC was measured on leaf sections obtained from the second fully developed upper leaves. They were quickly sealed within
plastic bags and fresh weights were determined immediately after
excision. After placing them in distilled water in test tubes for
4 h at room temperature (nearly 23 C) and under the low light
environment of the laboratory, turgid masses were estimated. Dry
masses were measured after drying the leaf samples in oven for
48 h at 70 C. Finally, RWC was calculated by the Eq. (5) (Smart and
Bingham, 1974):
RWC(%) = (
FreshWeight DryWeight
) 100
TurgidWeight DryWeight
(5)
Free proline content in the leaves was measured using the

method of Bates et al. (1973). 500 mg of fresh mature leaves were
crushed in 10 ml of 3% aqueous sulphosalycylic acid and the extract
was ltered. 2 ml of the extract was added into the test tube containing 2 ml of ninhydrin reagent and 2 ml of glacial acetic acid.
The reaction mixture was heated in a boiling water bath at 100 C
for one hour. After cooling the mixture on ice, 4 ml of toluene was
added and thoroughly mixed. Finally, the toluene phase was separated and its absorbance measured at 520 nm using a HITACHI
U1800 spectrophotometer against toluene blank.
Total leaf soluble sugars were measured using the method of
Irigoyen et al. (1992). 500 mg of dried leaves was grinded in 5 ml
of 95% ethanol. Then, 0.1 ml of homogenized extract was mixed
with 3 ml of anthrone (150 mg anthrone, 100 ml of 72% sulphuric
acid). The reaction extract was placed in a boiling water bath for
10 min. The light absorption of the samples was measured spectrophotometrically at 625 nm. Total leaf soluble sugars content was
determined using glucose standard curve.
The content of chlorophyll in fresh leaves was measured
spectrophotometrically using the method of Lichtenthaler and
Buschmann (2001). One gram of fresh leaf tissue was crushed using
mortar and pestle containing 10 ml of acetone (80%). The light
absorption of leaf extract solution was recorded at 662 and 645 nm.
Finally the results were reported as mg of pigment per g of leaf fresh
weight.
2.4. Measurement of plant dry mass, seed yield and yield
attributes
At 70% physiological maturity, plant height, number of umbels
per plant and number of fruits per umbel of ve plants per experimental unit were measured. The harvested plants air dried for 7
days and subsequently, 1000-seed weight were determined. Seed
yield and above ground dry mass (DM) were determined from the
central 2.25 m2 portion in each plot and expressed as g m2 . The
DM was determined after the samples were oven-dried for 72 h at
70 C.
2.5. Measurement of seed essential oil content
At the 70% physiological maturity stage the fennel seeds were
harvested, air dried for 7 days and subsequently, hydrodistillation of essential oil was done on a crushed 100 g sample of each
experimental unit according to Clevenger (1928) by the Clevengers
apparatus (Borosil, India). 500 ml of deionized water was added to
497
100 g of grinded seeds. Distillation process continued for 4 h, boiling

at 100 C. The essential oil phase was separated, dried over anhydrous sodium sulfate, and conserved in a dark glass bottle at 4 C
for further analysis.
2.6. Determination of stress susceptibility index
The stress susceptibility index (SSI) was calculated according to
Eqs. (6) and (7) (Fischer and Maurer, 1978):
SSI =
1 Ysg /Yng
SI
SI = 1
Ys
Yn
(6)
(7)
where Ysg is the yield of genotype under stress conditions, Yng is the
yield of genotype under non-stress conditions,Ys is the mean yield
of all genotypes under stress conditions, and Yn is the mean yield of
all genotypes under non-stress conditions. A lower SSI represents
a higher drought tolerance.
2.7. Determination of irrigation water use efciency
Irrigation water use efciency (IWUE) is specied as the ratio of
the crop yield to irrigation water applied (Howell, 1994). Irrigation
water use efciencies for seed yield (IWUESY ), dry mass (IWUEDM )
and oil yield (IWUEOY ) were determined by dividing seed yield
(g m2 ), shoot dry mass (g m2 ) and oil yield (ml m2 ) to total water
(m3 ) applied to m2 for each irrigation level, respectively.
2.8. Statistical analysis
Analysis of variance was done to detect differences among the
treatments using the general linear model (GLM) in SAS software
(SAS Institute, 1999). Least signicant difference (LSD, P 0.05) test
was employed to separate the means, where data were found statistically signicant at P 0.05.
3. Results
All studied traits of fennel in this study, including plant DM
and height, number of umbels per plant, fruits per umbel and
seeds per m2 , 1000-seed weight, seed yield, harvest index, seed
essential oil content and yield, irrigation water use efciencies
for seed yield (IWUESY ), shoot dry mass (IWUEDM ) and oil yield
(IWUEOY ), leaf water potential and relative water content, leaf proline and soluble sugars contents, leaf chl a, chl b, chl a + b and chla/b
were signicantly (P 0.05) affected by soil moisture, genotype and
the interaction effects of soil moisture genotype (Table 3). Main
means for the above traits are briey explained rst, and then the
interaction means are dealt with in more details afterward.
Averaged over genotypes, mean DM and plant height of fennel
were decreased by 65% and 34%, respectively, when grown under
severe drought, compared to control (Table 4). Genotype Mashhad produced the greatest mean DM and Ardabil the smallest,
averaged over the irrigation regimes. Genotypes Isfahan and Avicenna indicated the greatest plant height and Ardabil the smallest,
averaged over the irrigation regimes. Furthermore, mean number
of umbels per plant and fruits per umbel were decreased by 71% and
59%, respectively, when subjected to the severe drought, compared
to control (Table 5). In the same way with the latter attributes, mean
number of seeds per m2 and 1000-seed weight were decreased
by 50% and 24%, respectively (Tables 6 and 7). Genotype Yazd
produced the greatest mean number of umbel and Kerman the
smallest, averaged over the irrigation regimes. Genotype Shiraz
produced the greatest number of fruits per umbel and Kerman
498
Table 3
Analysis of variance for the effect of four irrigation levels on studied traits of fennel genotypes.
Traits
Df
Shoot dry mass
Plant height
Number of umbels per plant
Number of fruits per umbel
Number of seeds per m2
Seed yield
Harvest index
1000-seed weight
Seed essential oil content
Seed essential oil yield
Irrigation water use efciency of seed yield
Irrigation water use efciency of essential oil yield
Irrigation water use efciency of shoot dry mass
Leaf water potential
Relative water content (RWC)
Leaf proline content
Leaf soluble sugars content
Chlorophyll a content
Chlorophyll b content
Chlorophyll a + b content
Chlorophyll a/b
Mean of squares
Replications (R)
Irrigation levels (I)
RI
Genotypes (G)
IG
Error
2
222353.40**
37.63
10777.27**
66.512**
66990566
141.92
0.0425**
0.0989**
3.2621**
2.7036**
184.29
2.832**
237526.0**
3.0524**
13.2880**
16.668**
12.2937**
0.04365**
0.00068**
0.05427**
1.4574**
3
1585859.38**
8500.11**
1418975.55**
787.826**
3363967649**
58226.86**
0.0114**
4.6743**
0.134**
21.9403**
13501.95**
4.544**
362354.0**
6.6413**
351.3200**
382.507**
6.8180**
0.21674**
0.03437**
0.42301**
0.0670**
6
17040.46
10.83
3171.29
4.185
10925347
104.31
0.0010
0.0106
0.0007
0.1970
139.18
0.139
5728.0
0.0565
0.0427
5.626
0.2340
0.00138
0.00002
0.00146
0.0429
11
72710.53**
1795.92**
333187.04**
52.746**
436675008**
4237.64**
0.0138**
1.1605**
0.265**
2.8224**
5141.26**
3.321**
69780.5**
1.0652**
98.5280**
532.739**
2.8180**
0.07382**
0.01167**
0.14289**
0.0808**
33
18058.99**
217.57**
25951.56**
5.903**
82280120**
1035.89**
0.0024*
0.0289**
0.0289**
0.5435**
1009.27**
0.540**
14082.2**
0.0366**
11.3190**
38.107**
0.1323**
0.00952**
0.00151**
0.01788**
0.0498**
88
249.72
22.22
1256.50
0.729
24326136
208.65
0.0013
0.0086
0.00053
0.1106
268.04
0.132
206.0
0.0011
0.0013
0.253
0.0073
0.00003
0.00001
0.00004
0.0001
*, ** Signicant at the 5 and 1% levels of probability, respectively; ns, non signicant; df, degrees of freedom.
the smallest, when averaged over the irrigation regimes. Genotype

Shiraz produced the greatest number of seeds per m2 and Ardabil
the smallest, when averaged over the irrigation regimes. Genotype
Ardabil produced the greatest 1000-seed weight and Kerman the
smallest, when averaged over the irrigation regimes. In agreement
to the effects observed for the fennel yield attributes, mean seed
yield was decreased by 61% under severe drought, compared to control (Table 7). In contrary, mean harvest index was increased by 13%
under severe drought, compared to control (Table 7), though, the
greatest increase in harvest index (21%) was obtained under mild
drought. Genotypes Shiraz and Yazd produced the greatest seed
yield per plant and Kerman and Ardabil the smallest, averaged
over the irrigation regimes. Genotypes Urmia, Yazd, Kashan and
Isfahan produced the greatest harvest indices and Kerman the
smallest, when averaged over the irrigation regimes.
While mean seed essential oil content was increased by 6%
under mild drought, 7% under moderate drought and 5% under
severe drought, mean seed essential oil yield was decreased substantially (i.e. 58%) under severe drought, compared to control
(Table 8). Genotype Urmia indicated the greatest mean seed essential oil content (2.24%) and Ardabil the smallest (1.76%), averaged
over irrigation regimes. Genotypes Shiraz and Yazd produced the
greatest mean seed essential oil yield and Ardabil and Kerman the
smallest, averaged over irrigation regimes.
Mean IWUESY (Table 9), IWUEDM (Table 9), and IWUEOY
(Table 10) were decreased by 30%, 38% and 24%, respectively,
when fennel was subjected to severe drought, compared to control. Though, mean IWUESY remained unchanged, IWUEDM was
decreased by 16% and IWUEOY increased by 7% at the presence of
mild drought. Genotypes Shiraz and Yazd indicated the greatest
and Kerman and Ardabil the smallest mean IWUESY , averaged
over the irrigation regimes. Genotype Mashhad indicated the
greatest and Hamadan and Kerman the smallest mean IWUEDM ,
averaged over irrigation regimes. Genotypes Shiraz and Yazd
indicated the greatest and Ardabil and Kerman the smallest mean
IWUEOY , averaged over the irrigation regimes.
Mean leaf water potential and relative water content were
decreased by 48% and 9%, respectively, when fennel was grown
under severe drought compared to control (Table 11). Genotype
Yazd indicated the greatest mean leaf water potential (1.85 MPa)
and Birjand and Hamadan the smallest (2.77 and 2.75 MPa,
respectively), averaged over the irrigation regimes. Genotypes
Yazd and Kashan indicated the greatest mean leaf water content
(80%), Hamadan and Birjand the smallest (72.8%), averaged over
the irrigation regimes.
Mean leaf proline and soluble sugars contents were increased by
52% and 35%, respectively, when fennel was grown under severe
drought compared to control (Table 12). Genotype Shiraz indicated the greatest mean leaf proline and Ardabil the smallest,
averaged over the irrigation regimes. Genotype Shiraz indicated
the greatest mean soluble sugars content and Hamadan the smallest, averaged over the irrigation regimes.
Mean leaf chl a, chl b (Table 13) and chl a + b contents (Table 14)
were decreased by 27%, 28% and 28%, respectively, when fennel
was grown under severe drought compared to control. Genotype
Shiraz indicated the greatest mean leaf chl a and chl a + b and
Birjand and Kerman the greatest chl b and chla/b, respectively.
Genotype Ardabil indicated the smallest mean chl a, chl b and chl
a + b and Kashan the smallest chl a/b, averaged over the irrigation
regimes.
Plant DM, plant height, number of umbels per plant, number of
fruits per umbel, number of seeds per m2 , 1000-seed weight, seed
yield per m2 , seed essential oil yield, IWUESY , IWUEDM , IWUEOY ,
leaf water potential, leaf relative water, chl a, chl b and chl a+b
contents were invariably decreased in all fennel genotypes with
progressive drought but the extents of these decreases differed
between genotypes, leading to the statistically signicant genotype
irrigation interactions for all the above traits. The greatest and
smallest decreases in DM were observed in genotypes Avicenna
(76%), and Kerman (41%) but those in plant height were observed
in Birjand (58%) and Ardabil (15%), respectively (Table 4). The
greatest and smallest decreases in number of umbels per plant
were observed in genotypes Avicenna (89%) and Ardabil (89%),
and Kerman (46%), though those in number of fruits per umbel
were found to be in Hamadan (73%) and Yazd (40%), respectively
(Table 5). The greatest and smallest decreases in number of seeds
per m2 were observed in genotypes Avicenna (66%) and Yazd
(13%), respectively (Table 6). The greatest and smallest decreases
in 1000-seed weight were detected in Hamadan (31%) and Birjand (29%), and Yazd (16%) (Table 7), and those in seed yield
Table 4
Effects of irrigation level (I1, I2, I3 and I4 = 35, 55, 75 and 85% depletion of available soil water, respectively), genotype (G) and their interaction on shoot dry mass and plant height of fennel.
Shoot dry mass (g m2 )
Plant height (cm)
I1
I2
I3
I4
Avg.
I1
I2
I3
I4
Avg.
Urmia
Hamadan
Kerman
Shiraz
Birjand
Yazd
Ardabil
Avicenna
Kashan
Mashhad
Bushehr
Isfahan
Avg.
681.71dea
576.52fg
490.38hi
807.82c
706.35d
627.05ef
493.61hi
962.61a
816.85c
895.3b
929.29ab
829.21c
734.72A
397.07jk(42)b
462.06in(20)
397.34jk(19)
563.98g(30)
527.53gh(25)
567.22g(10)
347.47k-n(30)
538.77gh(44)
465.67i(-43)
661.59de(26)
577.99fg(38)
535.51gh(35)
503.52B
311.36m-p(54)
258.75p-t(55)
358.07klm(27)
439.03ij(46)
335.98lmn(52)
444.38ij(29)
252.59 p-u(49)
330.41lmn(66)
320.52l-o(61)
401.04jk(55)
371.06kl(60)
306.39m-q(63)
344.13C
216.88s-v(68)
199.77uv(65)
289.56n-r(41)
374.89kl(54)
200.64tuv(72)
304.84m-q(51)
179.54v(64)
233r-v(76)
252.25q-u(69)
325.41l-o(64)
267.92o-s(71)
234.79r-v(72)
256.62D
401.755Gc
374.276H
383.839H
546.429B
442.622F
485.872D
318.302I
516.198C
463.821E
570.836A
536.564B
476.478DE
96.66gh
115.83bc
75.00m-q
110.00bcd
108.33cde
92.50hi
57.50tu
126.67a
101.67efg
100.83efg
103.33d-g
117.50b
100.49A
87.50ij(9)
105.83efd(9)
67.50qrs(10)
92.50hi(16)
85.83ijk(21)
91.66hi(1)
61.66stu(+7)
101.67efg(20)
99.16fgh(2)
101.67efg(+1)
99.16fgh(4)
100.83efg(14)
91.25B
82.50j-m(15)
79.16k-n(32)
59.16tu(21)
85.83ijk(22)
75.00m-q(31)
75.83l-p(18)
55.83uv(3)
77.50l-o(39)
82.50j-m(19)
80.83j-n(20)
69.16p-s(33)
95.83gh(18)
76.59C
78.33k-n(19)
58.33tu(50)
55.83uv(26)
83.33jkl(24)
45.83w(58)
62.50r-u(32)
49.16vw(15)
70.00o-r(45)
78.33k-n(23)
70.00o-r(31)
65.00rst(37)
73.33n-q(38)
65.83D
86.25EF
89.79CDE
64.37I
92.91BC
78.75H
80.62GH
56.04J
93.95AB
90.41BCD
88.33DE
84.16FG
96.87A
a
b
c
Means in columns and rows (interaction) for each trait followed by the same lowercase letter(s) are not signicantly different at the 5% probability level.
Values in the parentheses are the percentage decrease () or increase (+) compared with the control (I1 ).
Means in each column (main effect of genotype) or row (main effect of irrigation level) for each trait followed by the same uppercase letter(s) are not signicantly different at the 5% probability level.
Table 5
Effects of irrigation level (I1, I2, I3 and I4 = 35, 55, 75 and 85% depletion of available soil water, respectively), genotype (G) and their interaction on number of umbels per plant and number of fruits per umbel of fennel.
Number of umbels per plant
Number of fruits per umbel
Genotypes
I1
I2
I3
I4
Avg.
I1
I2
I3
I4
Avg.
Urmia
Hamadan
Kerman
Shiraz
Birjand
Yazd
Ardabil
Avicenna
Kashan
Mashhad
Bushehr
Isfahan
Avg.
496.33gha
465.67g-j
175.83qrs
915.00bc
411.67jkl
845.67d
583.17ef
937.17ab
524.50fg
609.00e
615.50e
558.17ef
594.81A
409.00jkl(18)b
323.17mn(31)
244.67op(+39)
856.00cd(6)
345.00m(16)
979.67a(+16)
374.33lm(36)
488.17ghi(48)
435.00jkl(17)
604.00e(1)
455.33hij(26)
580.83ef(+4)
507.93B
281.50no(-43)
111.50t-w(76)
143.33r-u(18)
582.17ef(36)
119.50s-v(71)
614.50e(27)
142.83r-u(76)
182.33qr(81)
264.00nop(50)
318.33mn(48)
232.33opq(62)
248.67op(55)
270.08C
181.67qr(-63)
64.67vw(86)
95.00uvw(46)
377.33klm(59)
58.67w(86)
408.33jkl(52)
61.67vw(89)
101.17t-w(89)
137.17r-u(74)
220.67pq(64)
158.67qr(74)
183.67rst(67)
170.72D
342.13Ec
241.25G
164.71H
682.63B
233.71G
712.04A
290.50F
427.21C
340.17E
438.00C
365.46DE
392.83D
19.00bc
21.33a
11.00pqr
21.33a
16.33f-i
18.66bcd
18.00b-e
21.66a
17.00e-h
19.50b
15.66hij
19.00bc
18.20A
16.50e-i(13)
17.66c-g(17)
10.16rs(8)
17.83c-f(16)
11.33o-r(31)
17.33d-g(7)
15.66hij(13)
16.83e-i(22)
15.66hij(8)
16.50e-i(15)
13.83klm(12)
16.16g-j(15)
15.45B
12.50m-p(34)
12.66mno(41)
8.00t(27)
15.33ijk(28)
8.66st(47)
14.66jkl(21)
10.33qr(43)
11.00pqr(49)
11.83n-q(30)
12.50m-p(36)
10.66qr(32)
13.16lmn(31)
11.77C
7.83tu(59)
5.83w(73)
6.33uvw(42)
11.00pqr(48)
5.66w(65)
11.16o-r(40)
5.50w(69)
6.00vw(72)
7.50tuv(56)
7.66tu(61)
6.33uvw(60)
8.16t(57)
7.41D
13.95C
14.37C
8.87G
16.37A
10.50F
15.45B
12.37D
13.87C
13.00D
14.04C
11.62E
14.12C
a
b
c
Genotypes
499
500
Table 6
Effects of irrigation level (I1, I2, I3 and I4 = 35, 55, 75 and 85% depletion of available soil water, respectively), genotype (G) and their interaction on number of seeds per m2 and seed yield of fennel.
Seed yield (g m2 )
Number of seeds per m2

I1
I2
I3
I4
Avg.
I1
I2
I3
I4
Avg.
Urmia
Hamadan
Kerman
Shiraz
Birjand
Yazd
Ardabil
Avicenna
Kashan
Mashhad
Bushehr
Isfahan
Avg.
45087b-da
39393e-i
23885n-r
50596ab
42373c-f
37367e-j
25338m-q
54063a
44260b-e
34340g-k
41643d-g
49554abc
40658A
36125f-j(20)b
33704h-l(15)
32022i-m(+35)
38715e-i(24)
38553e-j(10)
48162a-d(+29)
17008rst(33)
27531k-p(50)
43149b-f(3)
40427d-h(+18)
31882i-m(24)
37345e-j(25)
35385B
23655n-r(48)
18107qrs(55)
17447rs(27)
32092i-m(37)
25953l-q(39)
30759j-n(18)
15118st(41)
22476o-s(59)
28082k-o(37)
20148p-s(42)
23361n-r(44)
20184p-s(60)
23115C
16986rst(63)
15184st(62)
18175qrs(24)
34003g-k(33)
16641rst(61)
32724h-m(13)
9159t(-64)
18443qrs(66)
22859o-s(49)
20276o-s(41)
19965p-s(53)
21515o-s(57)
20494D
30463DEc
26597EF
22882F
38851A
30880CD
37253AB
16656G
30628CD
34587BC
28798DE
29213DE
32150CD
160.00bcd
138.75d-g
66.19p-u
166.69abc
139.06d-g
125.44ghi
108.19i-l
187.06a
149.38c-f
134.63e-h
154.69cde
181.81ab
142.65A
123.63ghi(23)
108.88ijk(22)
85.25l-q(+29)
123.56ghi(26)
115.13hij(17)
158.25cd(+26)
67.44p-t(38)
87.63k-p(53)
137.44d-h(8)
149.56c-f(+11)
109.31ijk(29)
128.38f-i(29)
116.20B
75.06m-s(53)
51.00t-y(63)
44.31u-y(33)
97.94j-m(41)
68.63o-t(51)
95.06j-n(24)
54.13s-y(50)
62.38q-v(67)
81.81m-r(45)
68.88o-t(49)
74.31n-s(52)
63.13q-v(65)
69.71C
57.06s-y(64)
36.69xy(74)
41.19v-y(38)
92.69j-n(44)
39.00wxy(72)
91.31k-o(27)
34.13y(68)
46.00t-y(75)
60.75r-w(59)
61.81r-w(54)
56.00s-y(64)
58.88r-x(68)
56.29D
103.93CD
83.82F
59.23G
120.21A
90.45EF
117.51AB
65.96G
95.76DE
107.34BCD
103.71CD
98.57CDE
108.04BC
a
b
c
Table 7
Effects of irrigation level (I1, I2, I3 and I4 = 35, 55, 75 and 85% depletion of available soil water, respectively), genotype (G) and their interaction on 1000-seed weight and harvest index of fennel.
1000-seed weight (g)
Harvest index
Genotypes
I1
I2
I3
I4
Avg.
I1
I2
I3
I4
Avg.
Urmia
Hamadan
Kerman
Shiraz
Birjand
Yazd
Ardabil
Avicenna
Kashan
Mashhad
Bushehr
Isfahan
Avg.
3.550d-ga
3.516e-h
2.783t-w
3.283j-m
3.283j-m
3.350i-l
4.283a
3.450f-i
3.366h-k
3.933b
3.700cd
3.650cde
3.512A
3.416g-j(4)b
3.233k-n(8)
2.683u-x(4)
3.183m-p(3)
2.983qrs(9)
3.283j-m(2)
3.966b(7)
3.183m-p(8)
3.200l-o(5)
3.716c(6)
3.433f-j(7)
3.416g-j(6)
3.308B
3.166m-p(11)
2.816tu(20)
2.533xy(9)
3.033pqr(8)
2.633wx(20)
3.083n-q(8)
3.583c-f(16)
2.750uvw(20)
2.916rst(13)
3.450f-i(12)
3.166m-p(14)
3.116n-q(15)
3.020C
2.833stu(20)
2.416zya(31)
2.266a(19)
2.716uvw(17)
2.333za(29)
2.800tuv(16)
3.116n-q(27)
2.466yz(29)
2.650vwx(21)
3.066o-r(22)
2.800tuv(24)
2.733uvw(25)
2.683D
3.241Cc
2.995EF
2.566H
3.054ED
2.808G
3.129D
3.737A
2.962F
3.033EF
3.541B
3.27C
3.229C
0.238d-k
0.244d-j
0.133o
0.212e-k
0.198h-n
0.206g-n
0.221e-l
0.200h-n
0.183k-o
0.153mno
0.173l-o
0.221e-l
0.198C
0.319a(+35)
0.241d-k(2)
0.214e-l(+62)
0.226e-l(+7)
0.218e-l(+11)
0.294a-d(+43)
0.203g-n(8)
0.169l-o(16)
0.314ab(+71)
0.237d-k(+56)
0.191i-o(+11)
0.251c-h(+14)
0.240A
0.251c-h(+6)
0.197h-n(20)
0.132o(1)
0.236d-k(+12)
0.210f-m(+7)
0.214e-k(+4)
0.245d-h(+11)
0.185j-o(-8)
0.266a-f(+45)
0.169l-o(+11)
0.199h-n(+15)
0.220e-l(1)
0.210BC
0.261a-g(+10)
0.183k-n(25)
0.147no(+11)
0.260b-g(+23)
0.197h-n(1)
0.305abc(+48)
0.198j-n(11)
0.196j-n(3)
0.269a-e(+47)
0.189i-o(+24)
0.206g-n(+20)
0.270a-e(+23)
0.223AB
0.267A
0.216CDE
0.156F
0.233BCD
0.206DE
0.255AB
0.217CDE
0.187E
0.258AB
0.187E
0.192E
0.241ABC
a
b
c
Genotypes
Table 8
Effects of irrigation level (I1 , I2 , I3 and I4 = 35, 55, 75 and 85% depletion of available soil water), genotype (G) and their interaction on seed essential oil content and yield of fennel.
Seed essential oil yield (ml m-2 )
Seed essential oil content (%)

I1
I2
I3
I4
Avg.
I1
I2
I3
I4
Avg.
Urmia
Hamadan
Kerman
Shiraz
Birjand
Yazd
Ardabil
Avicenna
Kashan
Mashhad
Bushehr
Isfahan
Avg.
2.065mnoa
2.000rst
1.820ab
2.020pqr
1.905xy
2.090lmn
1.760d
2.235efg
1.835ab
2.040opq
1.875yz
1.935vwx
1.965C
2.220e-h(+8)b
2.095lm(+5)
1.980stu(+9)
2.145jk(+6)
2.015qrs(+6)
2.200ghi(+5)
1.855z a(+5)
2.320bc(+4)
1.965tuv(+7)
2.205f-i(+8)
2.000rst(+7)
2.070mno(+7)
2.089A
2.290cd(+11)
2.005qrs(0)
2.055nop(+13)
2.240ef(+11)
1.925wx(+1)
2.200ghi(+5)
1.765cd(0)
2.205f-i(1)
2.035o-r(+11)
2.255de(+11)
2.040opq(+9)
2.170ij(+12)
2.098B
2.390a(+16)
1.800bc(10)
2.120kl(+16)
2.350b(+16)
1.845za(3)
2.230e-h(+7)
1.670e(5)
1.950uvw(13)
1.915wx(+4)
2.310c(+13)
1.915wx(+2)
2.195hi(+13)
2.057B
2.241Ac
1.975EF
1.9938E
2.188BC
1.922G
2.180C
1.762H
2.177C
1.937G
2.202B
1.957F
2.092D
3.375bc
2.789d-g
1.186p-u
3.427bc
2.642e-i
2.662e-i
1.883j-n
4.271a
2.751efg
2.779d-g
2.937cde
3.524b
2.852A
2.817def(17)
2.297d-k(18)
1.652l-p(+39)
2.698e-h(21)
2.328f-i(12)
3.522b(+32)
1.247o-t(34)
2.052j-m(52)
2.725efg(1)
3.325bcd(+20)
2.170h-i(26)
2.697e-h(23)
2.4612B
1.768k-o(48)
1.038q-v(63)
0.931s-v(-22)
2.240g-k(35)
1.327o-s(50)
2.121i-l(20)
0.956r-v(49)
1.492n-r(65)
1.668l-p(39)
1.552m-q(44)
1.551m-q(47)
1.373n-s(61)
1.501C
1.395n-s(59)
0.656uv(76)
0.896s-v(24)
2.157h-l(37)
0.722tuv(73)
2.088j-m(22)
0.569tuv(70)
0.925s-v(78)
1.168p-u(58)
1.430n-s(49)
1.075q-v(63)
1.310o-s(63)
1.199D
2.339BC
1.695E
1.166F
2.631A
1.755E
2.598AB
1.164F
2.185CD
2.078CD
2.271C
1.933DE
2.226C
a
b
c
Table 9
Effects of irrigation level (I1 , I2 , I3 and I4 = 35, 55, 75 and 85% depletion of available soil water, respectively), genotype (G) and their interaction on irrigation water use efciencies of seed yield and shoot dry mass of fennel.
Irrigation water use efciency of seed yield (g m3 )
Irrigation water use efciency of shoot dry mass (g m3 )
Genotypes
I1
I2
I3
I4
Avg.
I1
I2
I3
I4
Avg.
Urmia
Hamadan
Kerman
Shiraz
Birjand
Yazd
Ardabil
Avicenna
Kashan
Mashhad
Bushehr
Isfahan
Avg.
132.46b-ga
114.92f-k
54.59tu
138.97b-f
114.37f-k
104.28h-n
93.88i-o
154.37ab
124.55d-h
110.05g-m
126.69d-h
150.52a-d
118.30A
127.22d-h(4)b
119.83g-m(5)
89.45l-q(+62)
124.96d-h(11)
115.60e-j(+2)
165.29a(+59)
70.73o-u(25)
89.18m-q(43)
141.72a-e(+14)
153.48abc(+40)
112.85f-l(12)
128.22c-h(16)
119.04A
93.93i-o(29)
63.53q-u(45)
55.40tu(0)
122.34e-h(13)
85.00m-r(26)
118.61e-i(+15)
70.23o-u(26)
75.57o-u(51)
102.95h-m(18)
85.26m-r(23)
91.12j-o(29)
79.43n-t48)
86.94B
84.59m-r(36)
54.32tu(54)
61.13r-u(+11)
137.88b-f(1)
57.93stu(50)
135.56b-g(+31)
51.79u(45)
67.07p-u(57)
90.68j-p(28)
91.07j-p(18)
82.42n-s(36)
87.53l-q(42)
83.49B
109.54BCc
85.65E
65.14F
131.03A
93.22DE
130.93A
71.65F
96.54CDE
114.97B
109.96B
103.27BCD
111.42B
558.68cde
481.54hij
418.26m-p
665.64b
588.17c
517.71fgh
425.49l-o
786.2a
687.37b
762.73a
751.81a
691.11b
611.22A
407.2n-r(28)
462.68jkl(4)
419.73m-p(+1)
564.29cde(16)
536.85efg(9)
578.65cd(+12)
372.57rst(13)
537.31efg(32)
468.23ijk(32)
668.42b(13)
589.59c(22)
528.45efg(24)
511.16B
385.66p-s(31)
327.5uv(32)
467.44ijk(+12)
539.34efg(19)
412.69n-q(30)
554.3c-f(+8)
339.23tu(21)
416.75m-p(47)
396.55o-s(43)
505.51ghi(34)
472.00ijk(38)
377.16q-t(46)
432.84C
325.9uv(42)
296.84vw(39)
440.98k-n(+6)
547def(18)
297.4vw(50)
452.3k-m(13)
277.1w(35)
341.61tu(57)
366.79st(47)
482.67hij(37)
397.75o-s(48)
342.96tu(51)
380.78D
419.4H
392.1I
436.6G
579.1B
458.8F
525.7D
353.6J
520.5D
479.7E
604.8A
552.8C
484.9E
a
b
c
Genotypes
Means in columns and rows (interaction) for each trait followed by the same lowercase letter(s) are notsignicantly different at the 5% probability level.
Means in each column (main effect of genotype) or row (main effect of irrigation level) for each treatment followed by the same uppercase letter(s) are not signicantly different at the 5% probability level.
501
502
Table 10
Effects of irrigation level (I1 , I2 , I3 and I4 = 35, 55, 75 and 85% depletion of available soil water, respectively), genotype (G) and their interaction on irrigation water use efciency of seed essential oil yield of fennel.
Irrigation water use efciency of seed essential oil yield (ml m-3 )
Genotypes
I1
2.700c-g
2.299e-l
0.969u
2.786cde
2.167g-o
2.167g-o
1.613n-t
3.438ab
2.257e-l
2.254e-l
2.389d-j
2.905bcd
2.328A
2.813cde(+4)
2.305e-l(+5)
1.718l-s(+89)
2.684c-h(4)
2.330d-k(+10)
3.616a(+72)
1.294r-u(25)
2.049i-o(42)
2.772cde(+23)
3.407ab(+55)
2.225e-m(5)
2.678c-h(11)
2.490A
I3
I4
Avg.
2.169f-n(23)
1.284r-u(46)
1.152stu(+23)
2.758c-f(0)
1.639o-t(24)
2.614c-i(+24)
1.227r-u(25)
1.775k-r(50)
2.076i-o(10)
1.918j-p(14)
1.887j-q(22)
1.722l-s(42)
1.851B
2.036i-o(26)
0.972u(60)
1.318q-u(+45)
3.173abc(+15)
1.070tu(53)
3.063abc(+43)
0.858u(50)
1.338p-u(62)
1.736l-s(23)
2.105h-o(5)
1.578o-t(35)
1.933j-o(35)
1.764B
2.429Bc
1.715E
1.288F
2.850A
1.801DE
2.864A
1.248F
2.149BC
2.210BC
2.420B
2.019CD
2.309BC
Means in columns and rows (interaction) followed by the same lowercase letter(s) are not signicantly different at the 5% probability level.
Means in each column (main effect of genotype) or row (main effect of irrigation level) followed by the same uppercase letter(s) are not signicantly different at the 5% probability level.
Table 11
Effects of irrigation level (I1 , I2 , I3 and I4 = 35, 55, 75 and 85% depletion of available soil water, respectively), genotype (G) and their interaction on leaf water potential and RWC of fennel.
Leaf water potential (- MPa)
RWC (%)
Genotypes
I1
I2
I3
I4
Avg.
I1
I2
I3
I4
Avg.
Urmia
Hamadan
Kerman
Shiraz
Birjand
Yazd
Ardabil
Avicenna
Kashan
Mashhad
Bushehr
Isfahan
Avg.
1.498ta
2.117mno
2.067nop
1.810r
2.170lmn
1.445t
2.065nop
1.755rs
2.117mno
1.963qp
2.220klm
1.860gr
1.924D
1.808r(21)b
2.531ghi(20)
2.272kl(10)
2.013op(11)
2.582fgh(19)
1.653s(14)
2.480hi(20)
2.221klm(27)
2.427ij(15)
2.272kl(16)
2.531ghi(14)
2.169lmn(17)
2.246C
2.102no(40)
3.073bc(45)
2.641fg(28)
2.318jk(28)
3.073bc(42)
2.048op(42)
3.018cd(46)
2.802e(60)
2.803e(32)
2.641fg(35)
2.911de(31)
2.587fgh(39)
2.668B
2.555fgh(71)
3.268a(54)
2.645f(28)
2.501hi(38)
3.268a(51)
2.247kl(56)
3.218a(56)
3.1677ab(80)
2.962d(40)
2.809e(43)
3.015cd(36)
2.965cd(59)
2.885A
1.991Gc
2.747A
2.406E
2.161F
2.773A
1.848H
2.695B
2.486D
2.577C
2.421E
2.669B
2.395E
84.18a
76.20r
79.68g
81.18e
75.21u
84.17a
78.20l
82.19c
82.18c
82.18c
82.69b
81.19e
81.66d(3)
73.67x(3)
79.67g(0)
79.67g(2)
73.69xw(2)
80.65f(4)
74.67v(5)
76.66p(7)
79.16i(4)
75.67s(8)
78.69j(5)
78.20kl(4)
77.67B
77.28n(8)
71.81z(6)
77.79m(2)
78.30k(4)
71.81z(5)
76.77o(9)
72.29y(8)
71.83z(13)
78.79j(4)
73.78wx(10)
76.33q(8)
77.31n(5)
75.34C
73.76wx(12)
69.32b(9)
77.75m(2)
77.30n(5)
70.28a(7)
77.75m(8)
67.29c(14)
65.30d(21)
79.29h(4)
75.34t(8)
73.79w(11)
75.34t(7)
73.54D
79.22B
72.75J
78.72D
79.11C
72.75J
79.83A
73.11I
74.00H
79.86A
76.74G
77.88F
78.01E
a
b
c
Urmia
Hamadan
Kerman
Shiraz
Birjand
Yazd
Ardabil
Avicenna
Kashan
Mashhad
Bushehr
Isfahan
Avg.
b
I2
a
Table 12
Effects of irrigation level (I1 , I2 , I3 and I4 = 35, 55, 75 and 85% depletion of available soil water), genotype (G) and their interaction on leaf proline and leaf soluble sugars content of fennel.
Leaf proline content (moles g1 )
Leaf soluble sugars content (mg g1 )
I1
I2
I3
I4
Avg.
I1
I2
I3
I4
Avg.
Urmia
Hamadan
Kerman
Shiraz
Birjand
Yazd
Ardabil
Avicenna
Kashan
Mashhad
Bushehr
Isfahan
Avg.
13.267qrsa
13.317qrs
10.800v-y
20.133i
13.783pqr
18.667k
9.8667y
13.333qrs
12.750str
14.733nop
8.350z
12.717str
13.476C
15.750mn(+19)b
11.167u-x(17)
13.200qrs(+23)
34.600c(+72)
15.767mn(+15)
22.85gh(+23)
11.717tuv(+19)
10.150wxy(24)
17.283l(+36)
18.800jk(+28)
12.183stu(+46)
16.233lm(+28)
16.642B
19.433ijk(+47)
9.900y(26)
18.883ijk(+75)
43.067a(+114)
15.733mn(+15)
27.800e(+49)
9.9167xy(+1)
14.150opq(+7)
19.933ij(+57)
22.033gh(+50)
22.050gh(+165)
17.317l(+37)
20.018A
19.467ijk(+47)
11.333uvw(-15)
23.267g(+116)
41.067b(+104)
15.150mno(+10)
30.267d(+63)
8.600z(13)
11.317uvw(-16)
18.900ijk(+49)
24.900f(+70)
21.633h(+160)
19.483ijk(+54)
20.449A
16.979Dc
11.429I
16.538E
34.717A
15.108G
24.896B
10.025J
12.238H
17.217D
20.117C
16.054F
16.438EF
2.835u-y
2.415abc
2.52zab
3.36j-n
1.995e
2.94t-x
2.205cde
3.045q-v
2.31bcd
3.15n-t
2.73w-z
2.10de
2.633C
3.296k-p(+17)
2.614zya(+9)
2.841u-y(+13)
3.864de(+16)
2.728w-z(+37)
3.410i-m(+16)
2.728w-z(+24)
3.069q-u(+1)
2.955r-w(+28)
3.637e-i(+16)
3.182m-r(+17)
2.500zab(+20)
3.069B
3.802def(+35)
2.944s-x(+22)
3.312k-o(+32)
4.538ab(+36)
3.066p-u(+54)
3.925d(+34)
3.189m-r(+45)
3.434i-l(+13)
3.557g-j(+54)
4.170c(+33)
3.680e-h(+35)
2.821v-y(+35)
3.536A
3.606f-i(+28)
2.704xyz(+12)
3.735d-g(+49)
4.508ab(+35)
2.833u-y(+43)
4.379bc(+49)
2.962r-w(+35)
3.091o-t(+2)
3.348j-n(+45)
4.636a(+48)
3.477h-k(+28)
3.220l-q(+54)
3.542A
3.385D
2.669I
3.102FG
4.067A
2.655I
3.663C
2.771H
3.160F
3.042G
3.898B
3.267E
2.660I
a
b
c
Table 13
Effects of irrigation level (I1, I2, I3 and I4 = 35, 55, 75 and 85% depletion of available soil water, respectively), genotype (G) and their interaction on Chl a and chl b content of fennel.
Chl a (mg g1 )
Chl b (mg g1 )
Genotypes
I1
I2
I3
I4
Avg.
I1
I2
I3
I4
Avg.
Urmia
Hamadan
Kerman
Shiraz
Birjand
Yazd
Ardabil
Avicenna
Kashan
Mashhad
Bushehr
Isfahan
Avg.
0.588ija
0.547lmn
0.650fg
0.687e
0.840a
0.660f
0.449x
0.504rst
0.720bc
0.693de
0.598i
0.558kl
0.624A
0.553lm(6)b
0.524opq(5)
0.696de(+8)
0.730b(+7)
0.709cd(16)
0.571jk(14)
0.420z(7)
0.467w(8)
0.517pqr(29)
0.530nop(24)
0.510qr-s(15)
0.524opq(6)
0.563B
0.503rst(15)
0.500rst(9)
0.486tuv(26)
0.561kl(19)
0.531nop(37)
0.536mno(19)
0.373b(17)
0.396a(22)
0.496stu(32)
0.503rst(28)
0.447xy(26)
0.432xyz(23)
0.480C
0.508qrs(14)
0.430zy(22)
0.468w(28)
0.635gh(8)
0.420z(50)
0.620h(7)
c(26)
0.329c(35)
0.473vw(35)
0.482uvw(31)
0.345c(43)
0.416z(26)
0.455D
0.538Fc
0.500G
0.575D
0.654A
0.625B
0.596C
0.394K
0.424J
0.551E
0.552E
0.475I
0.483H
0.223m
0.223m
0.249h
0.274d
0a
0.263f
0.169z
0.198t
0.279c
0.259g
0.232k
0.222mn
0.2446A
0.210o(7)
0.202rs(10)
0.250h(+1)
0.265e(4)
0.293b(14)
0.203qr(23)
0.158cd(7)
0.182wx(9)
0.226l(19)
0.211o(19)
0.220n(5)
0.210o(6)
0.2195B
0.201s(11)
0.197t(12)
0.184vw(27)
0.232k(16)
0.223m(35)
0.203qr(23)
0.148e(13)
0.164a(18)
0.206p(27)
0.201s(23)
0.174y(25)
0.160b(28)
0.1914C
0.185v(17)
0.156d(30)
0.181x(28)
0.237j(14)
0.158c(54)
0.239i(9)
0.129h(24)
0.139g(30)
0.205qp(27)
0.191u(27)
0.125i(47)
0.145f(35)
0.1746D
0.205F
0.194G
0.216E
0.252B
0.253A
0.227D
0.151K
0.170J
0.229C
0.215E
0.188H
0.184I
a
b
c
Genotypes
503
0.811h
0.770jk
0.900e
0.961c
1.180a
0.923d
0.618r
0.702n
0.999b
0.953c
0.830g
0.780ij
0.869A
0.743G
0.695H
0.792D
0.906A
0.879B
0.824C
0.546K
0.595J
0.781E
0.768F
0.663I
0.667I
0.694no(15)
0.587s(24)
0.649q(28)
0.873f(10)
0.579st(51)
0.859f(7)
0.463w(25)
0.468w(34)
0.678op(33)
0.673p(30)
0.470w(44)
0.561t-u(-29)
0.630D
0.704n(14)
0.697n(10)
0.671p(26)
0.794hi(18)
0.754kl(37)
0.739lm(20)
0.522v(16)
0.560u(21)
0.702n(30)
0.704n(27)
0.622r(26)
0.593s(25)
0.672C
0.763jk(7)
0.726m(6)
0.947c(+6)
0.995b(+4)
1.002b(16)
0.774j(17)
0.579st(7)
0.649q(8)
0.743ml(26)
0.742ml(23)
0.731m(12)
0.734m(6)
0.782B
b
a
Urmia
Hamadan
Kerman
Shiraz
Birjand
Yazd
Ardabil
Avicenna
Kashan
Mashhad
Bushehr
Isfahan
Avg.
2.82bcd(+5)
2.843bc(+13)
2.673g-m(1)
2.763c-f(+7)
2.741d-h(+8)
2.670g-m(+4)
2.656h-n(4)
2.442tuv(7)
2.381uv(11)
2.606k-p(6)
2.848bc(+8)
2.969a(+15)
2.701A
2.583n-q(5)
2.612k-o(+4)
2.730e-i(+2)
2.494rst(4)
2.457tu(4)
2.721e-j(+6)
2.593l-p(6)
2.492rst(5)
2.480st(7)
2.583n-q(7)
2.642j-n(1)
2.772cde(+8)
2.597A
2.717e-j(+1)
2.676f-l(+6)
2.868b(+7)
2.843bc(+10)
2.501q-t(2)
2.893ab(+12)
2.747d-g(+1)
2.650i-n(+2)
2.358v(12)
2.587m-p(7)
2.386uv(11)
2.576n-r(1)
2.650A
2.707e-j
2.525p-t
2.681f-k
2.587m-p
2.546o-s
2.588m-p
2.740d-h
2.621k-o
2.656h-n
2.754d-g
2.658h-n
2.583n-q
2.637A
I4
I3
I2
I1
Chl a/b
Avg.
I4
I3
I2
I1
Genotypes
Chl a + b (mg g-1 )
Table 14
Effects of irrigation level (I1 , I2 , I3 and I4 = 35, 55, 75 and 85% depletion of available soil water, respectively), genotype (G) and their interaction on Chl a+b and Chl a/b of fennel.
2.708C
2.664E
2.738A
2.672E
2.561G
2.718B
2.684D
2.551H
2.469I
2.632F
2.633F
2.725B
Avg.
504
per m2 were observed in genotypes Avicenna (75%) and Yazd

(27%), respectively (Table 6). Harvest index of the majority of fennel
genotypes was increased under the severe drought, compared to
control, the maximum increases were observed in genotypes Yazd
and Kashan (Table 7). Though, harvest indices of Hamadan and
Ardabil were decreased and that of Birjand remained unchanged.
Seed essential oil content of all genotypes was increased at the
mild and moderate drought levels, with the exception of genotypes Hamadan, Ardabil and Avicenna, where they remained
unchanged at the moderate drought level, compared to control
(Table 8). The latter genotypes contrasted the remaining genotypes
as their essential oil contents decreased when subjected to the
severe drought level. The greatest increase (16%) in seed essential
oil content due to the severe drought was detected in genotypes
Urmia, Kerman and Shiraz. The greatest and smallest decreases
in seed essential oil yield were found in genotypes Avicenna (78%)
and Hamadan (76%), and Yazd (22%) and Kerman (24%), respectively (Table 8).
Water use efciency attributes responded to drought in a
genotype-specic manner. IWUESY for the majority of genotypes
was increased with mild drought but it decreased with severe
drought for all genotypes, with the exception of genotypes Yazd
and Kerman where it increased in comparison to control (Table 9).
IWUEDM was decreased in all genotypes under severe drought,
with the exception of genotype Kerman where it increased in
comparison to the control (Table 9). IWUEOY for the majority of
genotypes was increased with mild drought but it decreased with
severe drought for all genotypes, with the exception of genotypes
Yazd, Kerman and Shiraz where it increased in comparison to
control (Table 10). The greatest decreases in the water use efciency
attributes were in the range of 5762% and found to belong to genotype Avicenna, though, genotype Kerman indicated increases in
all latter attributes when it was subjected to severe drought.
The greatest (80%) and smallest (28%) decreases in leaf water
potential were observed in genotypes Avicenna and Kerman,
respectively and those in leaf relative water content were observed
in Avicenna (21%), and Kerman (2%) and Shiraz (5%), though,
the latter decreases were not statistically signicant (Table 11).
Leaf proline content of fennel genotypes indicated contrasting
responses to progressive drought, where it decreased in Hamadan,
Ardabil and Avicenna and increased in the remaining genotypes, with genotype Bushehr indicating the greatest increase
among all genotypes (Table 12). Leaf soluble sugars content was
increased in all fennel genotypes with progressive drought but the
extent of these increases differed between genotypes (Table 12).
The greatest (54%) and smallest (2%) increases in leaf soluble sugars content were observed in genotypes Isfahan and Avicenna,
respectively. The greatest decreases in leaf chl a, chl b and chl
a + b (50%, 54% and 51%) were observed in genotype Birjand and
the smallest decreases (7%, 9% and 7%) were found in genotype
Yazd (Tables 13 and 14). Leaf chl a/b of genotypes Kerman, Ardabil, Avicenna, Kashan and Mashhad tended to decrease with
drought, though, it increased in the remaining genotypes (Table 14).
4. Discussion
It appears from our data that plant growth and yield attributes
in all fennel genotypes tended to decrease with progressive water
decit. Drought depressed both primary (i.e. seed number per m2
and seed size) and secondary (i.e. number of umbels) yield components of this medicinal plant. In fact, number of umbels, number
of seeds per m2 and seed size were decreased to nearly 30%, 40%
and 75% of the control value, respectively, due to the imposition of
prolonged severe drought. Marked decreases in yield attributes at
present work agree with that of Mohamed and Abdu (2004), where
water decit led to decreases in plant height, branching and, therefore, seed and oil yield of fennel. Our ndings on fennel are in partial
agreement with those of Puppala et al. (2005) on Lesquerella fendleri, where pod per plant was decreased but seed per pod and seed
size were not affected by water decit. Variation in grain yield, yield
components and plant height and dry matter of other crop species
such as wheat (Zhang et al., 2005) with different irrigation schemes
has been postulated. It is generally accepted that yield components
are closely related to one another and do not inuence crop yields
independently. When a crop is subjected to prolonged drought, as
has been the case in the present study, the yield reductions could be
substantial and due primarily to decreases in seed number rather
than the seed weight. The prolonged drought leads to inhibition
of ower development, failure of embryo fertilization and abortion of zygotes (Fageria et al., 2006). Intense drought is proposed
to reduce number of seeds per unit area through decreases in fertility and mild drought often decreases the grain weight through
limiting assimilate supply to developing seeds (Giunta et al., 1993).
On the contrary to growth attributes and yield and yield components, though, seed essential oil content of a majority of fennel
genotypes either increased or remained unchanged with progressive drought (Table 8). Our results agree with those of Semiz et al.
(2012), where they found that fennel essential oil content was
increased from 3.5% under control to 3.8% under salt-stress conditions. Though, unlike the latter report, in the present study essential
oil content of seeds did not exceed 2.4% at its maximal value. Agronomic practices such as time of harvest (Stefanini et al., 2006) and
level of soil fertility (Mohamed and Abdu, 2004) may leave meaningful effects on seed essential oil content of fennel. It is, therefore,
reasonable to imagine that lack of agreement between previous
reports and our ndings with regard to the seed essential oil content may be, at least partially, related to differences in agronomic
practices, including harvesting time and soil fertilization level. Furthermore, the drought tolerant set of genotypes tended to contain
a greater essential oil in their seeds, compared to the sensitive
group. Our results further agree with the report of Mohamed and
Abdu (2004), where they found that fennel essential oil percent was
increased with drought. As Noreen and Ashraf (2010) have suggested, since the decrease in seed yield (i.e. due to water decit)
outweighed the increase in essential oil percent, the essential oil
yield was decreased across fennel genotypes (Table 8) when water
was withheld under the conditions of the present research.
Differences in harvest index are known to be related to differences in distribution of photoassimilates and differences in the
pattern of retention and/or remobilization of the photosynthates
(Gent and Kiyomoto, 1989). Achievement of greater harvest indices
under water stress conditions has been postulated in an array of
crop species including wheat (Davidson and Campbell, 1984) and
corn (Brown, 1986) and our ndings with fennels harvest index
(Table 7) are in accordance with the latter reports. It is, therefore,
reasonable to suggest that fennels behavior, in terms of distribution of photoassimilates, in response to water deprivation is not
different from the other crop species.
Irrigation water use efciency of seed yield and essential oil
yield were remained unchanged under mild water decit but they
were decreased under severe water decit. The ability of different plant species to utilize water efciently in the production of
dry matter varies greatly. In an early experiment conducted by
Green and Read (1983), corn required nearly 38% and 34% as much
moisture to produce a unit of dry matter as did wheat and sunowers, respectively. WUE variations in response to water availability
seem also to be species-dependent. 30%, 40% and 39% improvements in WUE for dry matter production due to water stress were
reported for corn, sunowers and wheat, respectively (Green and
Read, 1983). Wheat subjected to decit irrigation had 26% greater
WUE for grain yield production, relative to the control (Zhang et al.,
505
2005). Water use efciency of fennel was found to be maximal at

irrigation water salinities of up to 811 dS m1 (Semiz et al., 2012).
In contrary to the latter reports, however, the most favorable water
availability conditions were led to the highest WUE for seed and
dry matter yields of L. fendleri (Puppala et al., 2004). A decrease in
WUE of castor in drier years has been documented recently (Lima
et al., 2013). Our ndings with fennel are consistent with those of
Silva et al. (2009), where Aloe Vera (Aloe barbadensis Miller) plants
were proven less effective in water use for leaf biomass and gel
production, when subjected to extreme irrigation regimes, relative to intermediate regimes. Maximal water use efciencies for
whole plant dry matter in our experiment (611 g m3 ) were found
to be smaller than those reported by Semiz et al. (2012) (nearly
1.99 g l1 ). Though, our data for fennel seed yield WUE (ranging
from 83 to 118 g m3 ) were in accordance to those reported for castor (ranging from 20 to 150 g m3 ) (Lima et al., 2013). A decrease
in irrigation water use efciency for seed yield provides evidence
that no preferential utilization of photoassimilates by fennel seeds
occur during the prolonged drought.
Reduction in leaf water potential depresses key processes such
as leaf enlargement and photosynthesis (Good and Maclagan,
1993). The effect of water stress on tissue and organ development
is reected in altered patterns of plant growth and development.
Continuous exposure to water decit progressively decreases leaf
size over time due to decreases in both cell division and expansion. Water and selected solutes move down energy gradients from
the soil into the plant in response to osmotic potentials existing on
opposite sides of root membranes. Osmotic adjustment could make
a signicant contribution to sustaining the growth and function
of most plant species through maintaining a high water content
in their protoplasts. Osmotic adjustment helps to maintain turgor
and cell volume and it, therefore, is regarded as an important adaptive mechanism by which plants coup with stresses (Valkmar et al.,
1998). It has frequently been assumed that osmolytes and their
modications in plant tissues could be of potential use in explaining plants performance under adverse environmental conditions.
A variety of compatible solutes are induced in osmotic adjustment
that apart from contributing to the maintenance of cell turgor and
volume, they play several protective roles (Volkmar et al., 1998).
From the data in Tables 11 and 12 it is obvious that on the contrary
to invariable decreases in leaf water potential and RWC, both leaf
proline and leaf soluble sugar contents are increased in most of the
fennel genotypes when stricken by water decit. A signicant correlation between leaf water potential and RWC (Table 16) indicates
that with increase in leaf water potential, RWC of fennel genotypes
increases. Since the latter correlation was greater under drought,
compared to control, it is tempting to suggest that maintaining RWC
under water deprivation is more attributable to those cellular processes that increase leaf water potential. Signicant correlations
between leaf water potential and leaf proline content in one hand
and between RWC and leaf proline content on the other (Table 16)
are indicative of osmotic adjustment in the drought-stressed fennel
(as a greater proportional increase in the latter osmolyte is found
under water decit conditions). Data obtained with species such
as soybean (Porcel and Ruiz-Lozano, 2004) and mangrove grass
(Mohsenzadeh et al., 2006) conrms ours, in that high correlations between plant water potential and osmolytes, compatible
solutes, amino acids and soluble carbohydrates have been detected.
In mangrove grass (Aeluropus lagopoides), the drought-induced 20%
decrease in RWC was associated with increases in soluble sugars
and proline in this graminous species (Mohsenzadeh et al., 2006).
In soybean, the concentration of leaf proline and soluble carbohydrates of water-deprived plants was increased with decrease in leaf
water potential (Porcel and Ruiz-Lozano, 2004).
Prevalence of proline among osmolytes that are commonly
found in high concentrations when plants are exposed to drought
506
Table 15
Rankings of 12 fennel genotypes based on the stress susceptibility index (SSI) (1 = the highest and 12 = the lowest) under severe drought (irrigation after 85% depletion of available soil water) and moderate drought (irrigation
after 75% depletion of available soil water).
Severe water decit
Moderate water decit
SSI
Ranking
Group
Genotype
SSI
Ranking
Group
Yazd
Kerman
Shiraz
Mashhad
Kashan
Bushehr
Urmia
Isfahan
Ardabil
Birjand
Hamadan
Avicenna
0.45
0.62
0.73
0.89
0.98
1.05
1.06
1.12
1.13
1.19
1.22
1.25
12
11
10
9
8
7
6
5
4
3
2
1
Tolerant
Tolerant
Tolerant
Tolerant
Moderate
Moderate
Moderate
Moderate
Susceptible
Susceptible
Susceptible
Susceptible
Yazd
Kerman
Shiraz
Kashan
Mashhad
Ardabil
Birjand
Bushehr
Urmia
Hamadan
Isfahan
Avicenna
0.47
0.65
0.81
0.88
0.96
0.98
0.99
1.02
1.04
1.24
1.28
1.3
12
11
10
9
8
7
6
5
4
3
2
1
Tolerant
Tolerant
Tolerant
Tolerant
Moderate
Moderate
Moderate
Moderate
Susceptible
Susceptible
Susceptible
Susceptible
Table 16
Correlation coefcients among different traits of fennel genotypes.
Traits
10
11
12
13
14
15
16
17
18
19
20
21
22
1-Shoot dry mass

2-Plant height
3-Number of umbels per plant
4-Number of fruits per umbel
5-Number of seeds per m2
6-Seed yield
7-Harvest index
8-1000-seed weight
9-Seed essential oil content
10-Seed essential oil yield
11-IWUE of seed yielda
12-I WUE of shoot dry mass
13-IWUE of oil yield
14-Leaf water potential
15-Relative water content
16-Leaf proline content
17-Leaf soluble sugars content
18-Chl a content
19-Chl b content
20-Chl a + b content
21-Chl a/b
22-SSIb
1
0.65*
0.62*
0.39
0.68*
0.72**
0.06
0.03
0.58**
0.72**
0.69*
0.98**
0.71**
0.23
0.35
0.57*
0.65*
0.30
0.34
0.31
0.27
0.20
1
0.39
0.61*
0.74**
0.76**
0.33
0.11
0.61*
0.76**
0.67*
0.54
0.71**
0.18
0.22
0.30
0.22
0.16
0.21
0.17
0.30
0.20
1
0.79**
0.72**
0.79**
0.42
0.21
0.60*
0.82**
0.84**
0.65*
0.85**
0.69*
0.46
0.79**
0.74**
0.33
0.28
0.31
0.12
0.52
1
0.58*
0.75**
0.60*
0.36
0.59*
0.79**
0.75**
0.36
0.78**
0.52
0.20
0.52
0.49
0.08
0.05
0.07
0.08
0.11
1
0.91**
0.50
0.26
0.62*
0.89**
0.90**
0.64*
0.89**
0.53
0.54
0.72**
0.49
0.61*
0.64*
0.62*
0.21
0.33
1
0.62*
0.13
0.65*
0.96**
0.98**
0.67*
0.97**
0.54
0.51
0.66*
0.55
0.39
0.42
0.40
0.16
0.21
1
0.25
0.21
0.55
0.64*
0.12
0.57*
0.47
0.38
0.28
0.06
0.16
0.17
0.16
0.007
0.04
1
0.07
0.09
0.14
0.02
0.11
0.003
0.09
0.09
0.15
0.48
0.49
0.49
0.12
0.18
1
0.81**
0.64*
0.59*
0.78**
0.76**
0.46
0.55
0.68*
0.34
0.27
0.32
0.21
0.32
1
0.95**
0.68*
0.99
0.66*
0.50
0.67*
0.64*
0.39
0.38
0.39
0.06
0.25
1
0.67*
0.97**
0.61*
0.58*
0.74**
0.62*
0.45
0.46
0.45
0.10
0.34
1
0.69*
0.30
0.41
0.65*
0.73**
0.37
0.39
0.38
0.18
0.36
1
0.68*
0.55
0.73**
0.67*
0.45
0.44
0.44
0.03
0.33
1
0.70*
0.64*
0.65*
0.40
0.28
0.36
0.47
0.67*
1
0.62*
0.56
0.43
0.37
0.41
0.20
0.69*
1
0.80**
0.74**
0.68*
0.72**
0.18
0.69*
1
0.46
0.40
0.45
0.13
0.63*
1
0.97**
0.99**
0.02
0.59*
1
0.98**
0.17
0.48
1
0.03
0.56
1
0.41
*
**
a
b
Signicant at p < 0.05.

Signicant at p < 0.01.
Irrigation water use efciency (IWUE).
Stress susceptibility index (SSI) was calculated based on severe water decit (irrigation after 85% depletion of available soil water).
Genotype
and salt stresses is postulated (Porcel and Ruiz-Lozano, 2004). The

exact role of proline with regard to plants response to environmental stresses has been a focus of much research in recent decades,
though, still controversy exists on the nature of this role (Delauney
and Verma, 1993). Stabilizing proteins and enzymes and buffering the cellular redox potential (Demiral and Trkan, 2005) and
as a result, conferring enzyme protection and increasing membrane stability under various conditions (Nasir Khan et al., 2007)
have been proposed as main mechanisms through which proline
protects the cells from being harmed from stresses. Proline accumulation is known to be an outcome of proteolysis and decreases
in protein synthesis (Ashraf and Harris, 2004) and is hired in plant
as an adaptive advantage under water decit conditions (Delauney
and Verma, 1993). It, therefore, comes as no surprise that higher
RWC, leaf water potential, leaf proline and total leaf soluble sugars contents (Tables 11 and 12) were associated with greater DM
(Table 4) and seed yield (Table 6) under water decit conditions.
Since proline and (albeit in a lesser extent) total soluble sugars
accumulation in the fennel genotypes were correlated to their seed
yield, DM, RWC and leaf water potential (Table 16) it seems that
these osmolytes induce a partial relief from drought stress in fennel
at least with some of the genotypes used in this experiment. Genotypic variations for RWC and leaf water potential of fennel were
consistent with differences among the genotypes in key traits.
Based on the DM-based stress susceptibility index (SSI), fennel
genotypes were tentatively classied into drought tolerant (Yazd,
Kerman, Shiraz and Mashhad), moderately tolerant (Kashan,
Bushehr Urmia and Isfahan), and drought sensitive (Ardabil,
Birjand Hamadan and Avicenna) groups (Table 15). Drought tolerant group of genotypes were protected against drought, owing,
presumably, to the proportional higher leaf proline and total leaf
soluble carbohydrates. In fact, the genotypes Shiraz Yazd, Mashhad and Kerman ranked 1st to 4th, respectively, in terms of proline
accumulation (Table 12). However, the drought sensitive group had
lower leaf proline and total leaf soluble carbohydrates, compared to
the other groups. Osmotic adjustment is necessary to maintaining a
water potential gradient favorable to the water entrance from soil
into the roots and, therefore, fennel plants were able to achieve
this by raising the level of latter osmolytes. Osmotic adjustment
enabled maintaining higher leaf water potentials in genotypes Shiraz, Yazd, Mashhad and Kerman during drought and, perhaps,
kept the plants protected against oxidative stress and, therefore,
these cumulative effects increased the plant tolerance to moderate and severe drought levels. The drought sensitive group (i.e.
Ardabil, Birjand, Avicenna and Hamadan) had the lowest leaf
water potentials among the three groups. It is generally accepted
that RWC is positively correlated with plant tolerance to stresses
such as drought and salinity. In the present study the lowered RWC
in genotypes Ardabil, Avicenna, Birjand and Hamadan and the
greater RWC in genotypes Shiraz, Kerman and Yazd were associated with smaller and greater seed yields, respectively, in these
two sets of genotypes (Table 6).
Leaf chlorophyll concentration is directly related to the photosynthetic performance of the plants. In the present study, Chl a, Chl
b and, therefore, chl a + b of all fennel genotypes were decreased,
when they were deprived from water (Tables 13 and 14). The
decrease in leaf chlorophyll concentration could be due to increased
chlorophyllase activity and chlorophyll degradation, and an inhibition in the synthesis of photosynthetic pigments (Singh and Dubey,
1995). The extent of stress-induced decreases in leaf chl content is
expected to depend on the duration and severity of the drought
stress (Munne-Bosch and Alegre, 2000). Our ndings on fennel
confer with the latter expectation, as evidenced by a tendency of
chl attributes to decrease with drought intensication and prolongation. During the process of chlorophyll degradation Chl b is
converted into Chl, leading to increase in Chl a/b ratio with stress
507
conditions (Azizpour et al., 2010). Such indications were detected

in our experiment at least with a majority of the genotypes.
The signicant differences observed between fennel genotypes
in the osmolyte accumulation, growth attributes, photosynthetic
pigments, grain yield and attributes, essential oil content and water
use efciency in general and under water deprivation, in particular, suggest that there is much room for improving growth and
productivity of fennel by genetic manipulating these traits.
5. Conclusions
In the present study, decreases in leaf water potential, RWC
and chl content and plant growth characteristics were associated
with substantial decreases in grain yield of fennel at the presence
of severe drought. The negative effect of water stress on fennel
seed yield appeared to be mainly through seed number, rather
than seed weight. Fennel plants grown under mild water stress
produced marginally smaller seed and essential oil yields with comparable or greater WUE, compared to the plants grown under water
decit-free conditions. Considering the fact that adopting alternative crops with low irrigation needs is of high priority, adoption
of decit irrigation schemes to medicinal plants such as fennel will
allow the achievement of an improved WUE and may serve as a key
component in maintaining sustainable crop production in arid and
semi-arid environments. Osmolytes such as proline and, to a lesser
extent, total soluble sugars were found to make signicant contributions to osmoregulation in fennel. Unlike many reports on the
lack of correlation between proline and stress tolerance in different crop species, leaf water potential, DM and grain yield of fennel
genotypes responded to progressive drought, more or less, in proportion to their osmolytes content. From our ndings, fennel could
be appreciated as a promising species for semiarid climates, where
are characterized by varying degrees of water decit stress. Further studies utilizing the drought-tolerant set of fennel genotypes
described in this work will prove useful in elucidating the mechanisms of drought tolerance and potentiating alternative crops in
the face of the eminent challenge of water scarcity.
Acknowledgment
This study was supported with funds provided by the Isfahan
University of Technology, Isfahan, Iran.
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Industrial Crops and Products 76 (2015) 494508

Contents lists available at ScienceDirect

Industrial Crops and Products

Osmoregulation-mediated differential responses of eld-grown

Corresponding author. Fax +98 311 391 3453.

Water remains the single most important factor affecting world

E. Askari, P. Ehsanzadeh / Industrial Crops and Products 76 (2015) 494508

Organic C content (g kg1 )

Total N (mg kg1 )

Available P (mg kg1 )

Available K (mg kg1 )

Root zone depth (cm) 2011

Root zone depth (cm)

irrigation regimes were applied on early April and continued until

where rec is the recommended value for parsnip, and is the

E. Askari, P. Ehsanzadeh / Industrial Crops and Products 76 (2015) 494508

I1 (35% Depletion of ASW)

I2 (55% Depletion of ASW)

I1 (35% Depletion of ASW)

I2 (55% Depletion of ASW)

I3 (75% Depletion of ASW)

I4 (85% Depletion of ASW)

I3 (75% Depletion of ASW)

I4 (85% Depletion of ASW)

Cumulative applied water (mm)

mild, moderate and severe levels of drought stress, respectively. A

(%). Irrigation efciency was assumed to be 70% throughout the

E. Askari, P. Ehsanzadeh / Industrial Crops and Products 76 (2015) 494508

The mid-day leaf water potential was determined using a

Free proline content in the leaves was measured using the

100 g of grinded seeds. Distillation process continued for 4 h, boiling

E. Askari, P. Ehsanzadeh / Industrial Crops and Products 76 (2015) 494508

Irrigation levels (I)

the smallest, when averaged over the irrigation regimes. Genotype

Plant height (cm)

Number of fruits per umbel

E. Askari, P. Ehsanzadeh / Industrial Crops and Products 76 (2015) 494508

Number of seeds per m2

E. Askari, P. Ehsanzadeh / Industrial Crops and Products 76 (2015) 494508

Seed essential oil content (%)

Irrigation water use efciency of shoot dry mass (g m3 )

E. Askari, P. Ehsanzadeh / Industrial Crops and Products 76 (2015) 494508

E. Askari, P. Ehsanzadeh / Industrial Crops and Products 76 (2015) 494508

Leaf soluble sugars content (mg g1 )

E. Askari, P. Ehsanzadeh / Industrial Crops and Products 76 (2015) 494508

Chl a + b (mg g-1 )

E. Askari, P. Ehsanzadeh / Industrial Crops and Products 76 (2015) 494508

per m2 were observed in genotypes Avicenna (75%) and Yazd

E. Askari, P. Ehsanzadeh / Industrial Crops and Products 76 (2015) 494508

2005). Water use efciency of fennel was found to be maximal at

Moderate water decit

1-Shoot dry mass

Signicant at p < 0.05.

E. Askari, P. Ehsanzadeh / Industrial Crops and Products 76 (2015) 494508

E. Askari, P. Ehsanzadeh / Industrial Crops and Products 76 (2015) 494508

and salt stresses is postulated (Porcel and Ruiz-Lozano, 2004). The

conditions (Azizpour et al., 2010). Such indications were detected

E. Askari, P. Ehsanzadeh / Industrial Crops and Products 76 (2015) 494508

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