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Journal of Arid Land  2023, Vol. 15 Issue (5): 562-577    DOI: 10.1007/s40333-023-0011-x
Research article     
Morphological and physiological responses to drought stress of carob trees in Mediterranean ecosystems
Khouloud ZAGOUB, Khouloud KRICHEN, Mohamed CHAIEB, Lobna F MNIF*()
Laboratory of Ecosystems and Biodiversity in Arid Land of Tunisia, Faculty of Sciences, University of Sfax, Sfax 3000, Tunisia
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The greatest failure rate of reforestation programs is basically related to water deficit, especially at the seedling stage. Therefore, the main objective of this work is to investigate the responses of three accessions of carob trees (Ceratonia siliqua L.) with 2-year-old from different climate regions to drought generated by four water treatments: Tc (250 mm), T1 (180 mm), T2 (100 mm), and T3 (50 mm). The first accession (A1) comes from the protected national park of Ichkeul in northern Tunisia. This zone belongs to the bioclimatic sub-humid stage. The second accession (A2) comes from Melloulech, located in the center-east of Tunisia, belonging to the bioclimatic semi-arid stage. The third accession (A3) comes from the mountain of Matmata, located in the south of Tunisia, belonging to the bioclimatic hyper-arid stage. The experiment was undertaken in a greenhouse. Gaz exchange indices (net photosynthesis (A), stomatal conductance (gs), transpiration rate (E), and internal CO2 concentration (Ci)) were determined. Predawn (Ψpd) and midday (Ψmd) leaf water potentials, relative soil water content (SWC), and morphological parameters (plant height (H), number of leaves (NL), number of leaflets (Nl), and number of branches (NB)) were estimated. The results showed that significant differences (P<0.001) were found between physiological and morphological parameters of each accession. The highest growth potential was recorded for Tc treatment in both accessions A1 and A2. Significant decreases in gs, E, Ci, and SWC were recorded with the increases in water stress applied from treatment T1 to T3. Positive and significant correlations were found between SWC and Ψpd for all studied accessions. Ψpd and Ψmd decreased as water stress increased, ranging from -0.96 to -1.50 MPa at sunrise and from -1.94 to -2.83 MPa at midday, respectively, under control and T3 treatments. C. siliqua accessions responded to drought through exhibiting significant changes in their physiological and morphological behavior. Both accessions A1 and A2 showed greater drought tolerance than accession A3. These seedlings exhibit different adaptive mechanisms such as stress avoidance, which are aimed at reducing transpiration, limiting leaf growth, and increasing root growth to exploit more soil water. Therefore, C. siliqua can be recommended for the ecological restoration in Mediterranean ecosystems.

Key wordsCeratonia siliqua L.      water stress      soil moisture      stomatal conductance      predawn leaf water potential      minimum water potential     
Received: 25 October 2022      Published: 31 May 2023
Corresponding Authors: *Lobna F MNIF (E-mail address:
Cite this article:

Khouloud ZAGOUB, Khouloud KRICHEN, Mohamed CHAIEB, Lobna F MNIF. Morphological and physiological responses to drought stress of carob trees in Mediterranean ecosystems. Journal of Arid Land, 2023, 15(5): 562-577.

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Accession Code Longitude Latitude Mean annual precipitation (mm) Type of climate
(Emberger, 1954)
Ichkeul A1 09°40°14''E 37°08′24''N 600 Sub-humid
Melloulech A2 10°59°96''E 35°11°93''N 300 Semi-arid
Matmata A3 10°05°30''E 33°32°32''N 200 Hyper-arid
Table 1 Geographical origin and environmental conditions of the three accessions of Ceratonia siliqua
Fig. 1 Relative humidity and monthly mean temperature in the greenhouse during experimental period in 2020
Code Water treatment (mm) Explanation of water treatments
Tc 250 This is average precipitation of ideal year under arid region of Tunisia, which has a probability of occurrence of 0.2% according to Floret et al. (1981).
T1 180 This is usual average precipitation in the arid region of Tunisia according to Le Houerou (1969).
T2 100 This is usual average precipitation of the upper Saharan bioclimate according to IPCC (2021).
T3 50 This is usual average precipitation of the lower Saharan bioclimate of Tunisia.
Table 2 Split irrigation amounts applied during experimental period
Physiological parameter Morphological parameter
Net CO2 assimilation rate (A, µmol/(m2•s)) Plant height (H, cm)
Stomatal conductance (gs, mol H2O/m2•s)) Number of leaves formed (NL)
Transpiration rate (E, mmol H2O/(m2•s)) Number of leaflets formed (Nl)
Internal CO2 concentration (Ci, µmol/mol) Number of branching formed (NB)
Predawn leaf water potential (Ψpd, MPa) Chlorophyll content (SPAD value)
Midday leaf water potential (Ψmd, MPa)
Table 3 Parameters measured during experimental period
Index Source df F P
Plant height Accession 2 35.238 ***
Treatment 3 27.641 ***
Date 29 1.472 *
Accession×Treatment 6 7.900 ***
Accession×Date 18 0.042 ns
Treatment×Date 27 0.047 ns
Accession×Treatment×Date 51 0.030 ns
Number of leaves Accession 2 18.019 ***
Treatment 3 53.743 ***
Date 29 6.026 ***
Accession×Treatment 6 7.443 ***
Accession×Date 18 0.470 ns
Treatment×Date 27 0.483 ns
Accession×Treatment×Date 51 0.408 ns
Number of leaflets Accession 2 32.593 ***
Treatment 3 30.410 ***
Date 29 5.952 ***
Accession×Treatment 6 5.850 ***
Accession×Date 18 0.766 ns
Treatment×Date 27 0.460 ns
Accession×Treatment×Date 51 0.191 ns
Number of branches Accession 2 8.474 ***
Treatment 3 41.877 ***
Date 28 1.535 *
Accession×Treatment 6 6.002 ***
Accession×Date 18 0.329 ns
Treatment×Date 27 0.086 ns
Accession×Treatment×Date 51 0.103 ns
Chlorophyll content Accession 2 9.704 ***
Treatment 3 5.949 ***
Date 9 5.819 ***
Accession×Treatment 6 0.929 ns
Accession×Date 14 2.516 ***
Treatment×Date 21 3.313 ***
Accession×Treatment×Date 42 1.475 *
Table 4 Two-factor analysis of variation (ANOVA) results for morphological traits of Ceratonia siliqua
Fig. 2 Variation in morphological traits among Ceratonia siliqua accessions under different water treatments. (a), plant height; (b), number of leaves per plant; (c), number of leaflets per plant; (d), chlorophyll content. Tc, 250 mm/a; T1, 180 mm/a; T2, 100 mm/a; T3, 50 mm/a. SPAD, single-photon avalanche detector. *, P<0.05 level.
Index Source df F P
A Accession 2 15.243 ***
Treatment 3 17.538 ***
Date 7 9.350 ***
Accession×Treatment 6 2.895 **
Accession×Date 14 4.972 ***
Treatment×Date 21 0.984 ns
Accession×Treatment×Date 42 1.563 *
gs Accession 2 23.337 ***
Treatment 3 28.457 ***
Date 7 9.041 ***
Accession×Treatment 6 2.269 *
Accession×Date 14 8.514 ***
Treatment×Date 21 3.309 ***
Accession×Treatment×Date 42 2.353 ***
E Accession 2 24.099 ***
Treatment 3 77.320 ***
Date 7 22.216 ***
Accession×Treatment 6 2.049 *
Accession×Date 14 10.992 ***
Treatment×Date 21 5.416 ***
Accession×Treatment×Date 42 6.684 ***
Ci Accession 2 12.475 ***
Treatment 3 31.481 ***
Date 7 34.769 ***
Accession×Treatment 6 3.831 ***
Accession×Date 14 8.940 ***
Treatment×Date 21 1.964 **
Accession×Treatment×Date 42 3.121 ***
Table 5 Two-factor analysis of variation (ANOVA) results for gas exchange parameters of Ceratonia siliqua
Fig. 3 Variation in gas exchange parameter and water potential among Ceratonia siliqua accessions under different water treatments. (a), Ψpd, predawn leaf water potential; (b), Ψmd, midday leaf water potential; (c), A, net CO2 assimilation rate; (d), gs, stomatal conductance; (e), E, transpiration rate; (f), Ci, internal CO2 concentration.
Index Source df F P
Ψpd Accession 2 19.792 ***
Treatment 3 113.513 ***
Date 7 88.768 ***
Accession×Treatment 6 4.439 ***
Accession×Date 14 3.391 ***
Treatment×Date 21 9.809 ***
Accession×Treatment×Date 42 1.961 ***
Ψmd Accession 2 3.582 *
Treatment 3 159.107 ***
Date 7 293.495 ***
Accession×Treatment 6 11.729 ***
Accession×Date 14 8.902 ***
Treatment×Date 21 13.709 ***
Accession×Treatment×Date 42 4.778 ***
SWC Accession 2 9.825 ***
Treatment 3 102.434 ***
Date 7 31.546 ***
Accession×Treatment 6 1.121 ns
Accession×date 14 0.803 ns
Treatment×Date 21 4.994 ***
Accession×Treatment×Date 42 1.491 *
Table 6 Two-factor analysis of variation (ANOVA) results for leaf water potential and SWC of Ceratonia siliqua
Fig. 4 Relationships among physiological, morphological, and SWC (soil water content) parameters of Ceratonia siliqua accessions (A1-A3). (a), relationship between predawn leaf water potential (Ψpd) and SWC; (b), relationship between stomatal conductance (gs) and SWC; (c), relationship between chlorophyll content and SWC. SPAD, single-photon avalanche detector. *, P<0.05 level; **, P<0.01 level.
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