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Enhancement of some citrus rootstocks tolerance to drought stress by vermicompost and foliar application of trehalose Cover

Enhancement of some citrus rootstocks tolerance to drought stress by vermicompost and foliar application of trehalose

Open Access
|Dec 2025

Figures & Tables

Figure 1.

The influence of diverse levels of drought (70% FC and 40% FC), V.CO and exogenous TH on CRH (cm) (A), Shoots number (B) and Stem diameter (mm) (C).Three replicates (n = 3) for each season and the means of two se asons (2023 and 2024) make up the data. Duncan’s multiple range test indicates a significant difference at p ≤ 0.05 between the mean values ± SE of each parameter followed by various alphabetical letters. CRH, citrus rootstock height; FC, field capacity; SE, standard error; TH, trehalose; V.CO, vermicompost.
The influence of diverse levels of drought (70% FC and 40% FC), V.CO and exogenous TH on CRH (cm) (A), Shoots number (B) and Stem diameter (mm) (C).Three replicates (n = 3) for each season and the means of two se asons (2023 and 2024) make up the data. Duncan’s multiple range test indicates a significant difference at p ≤ 0.05 between the mean values ± SE of each parameter followed by various alphabetical letters. CRH, citrus rootstock height; FC, field capacity; SE, standard error; TH, trehalose; V.CO, vermicompost.

Figure 2.

The influence of diverse levels of drought (70% FC and 40% FC), V.CO and exogenous TH on citrus rootstock leaves number (A), LT (mm) (B) and LA (cm2) (C). Three replicates (n = 3) for each season and the means of two seasons (2023 and 2024) make up the data. Duncan’s multiple range test indicates a significant difference at p ≤ 0.05 between the mean values ± SE of each parameter followed by various alphabetical letters. FC, field capacity; LA, leaf area; LT, leaf thickness; SE, standard error; TH, trehalose; V.CO, vermicompost.
The influence of diverse levels of drought (70% FC and 40% FC), V.CO and exogenous TH on citrus rootstock leaves number (A), LT (mm) (B) and LA (cm2) (C). Three replicates (n = 3) for each season and the means of two seasons (2023 and 2024) make up the data. Duncan’s multiple range test indicates a significant difference at p ≤ 0.05 between the mean values ± SE of each parameter followed by various alphabetical letters. FC, field capacity; LA, leaf area; LT, leaf thickness; SE, standard error; TH, trehalose; V.CO, vermicompost.

Figure 3.

The influence of diverse levels of drought (70% FC and 40% FC), V.CO and exogenous TH on optical quality (A) and RWC (%) (B). Three replicates (n = 3) for each season and the means of two seasons (2023 and 2024) make up the data. Duncan’s multiple range test indicates a significant difference at p ≤ 0.05 between the mean values ± SE of each parameter followed by various alphabetical letters. FC, field capacity; RWC, relative water content; SE, standard error; TH, trehalose; V.CO, vermicompost.
The influence of diverse levels of drought (70% FC and 40% FC), V.CO and exogenous TH on optical quality (A) and RWC (%) (B). Three replicates (n = 3) for each season and the means of two seasons (2023 and 2024) make up the data. Duncan’s multiple range test indicates a significant difference at p ≤ 0.05 between the mean values ± SE of each parameter followed by various alphabetical letters. FC, field capacity; RWC, relative water content; SE, standard error; TH, trehalose; V.CO, vermicompost.

Figure 4.

The influence of diverse levels of drought (70% FC and 40% FC), V.CO and exogenous TH on nitrogen (N) (mg · 100 g−1 DW) (A), phosphorus (P) (mg · 100 g−1 DW) (B) and potassium (K) (mg · 100 g−1 DW) (C). Three replicates (n = 3) for each season and the means of two seasons (2023 and 2024) make up the data. Duncan’s multiple range test indicates a significant difference at p ≤ 0.05 between the mean values ± SE of each parameter followed by various alphabetical letters. DW, dry weight; FC, field capacity; SE, standard error; TH, trehalose; V.CO, vermicompost.
The influence of diverse levels of drought (70% FC and 40% FC), V.CO and exogenous TH on nitrogen (N) (mg · 100 g−1 DW) (A), phosphorus (P) (mg · 100 g−1 DW) (B) and potassium (K) (mg · 100 g−1 DW) (C). Three replicates (n = 3) for each season and the means of two seasons (2023 and 2024) make up the data. Duncan’s multiple range test indicates a significant difference at p ≤ 0.05 between the mean values ± SE of each parameter followed by various alphabetical letters. DW, dry weight; FC, field capacity; SE, standard error; TH, trehalose; V.CO, vermicompost.

Figure 5.

The influence of diverse levels of drought (70% FC and 40% FC), V.CO and exogenous TH on chlorophyll a (μg · cm2 DW) (A), chlorophyll b (μg · cm2 DW) (B) and carotenoids (μg · cm2 DW) (C). Three replicates (n = 3) for each season and the means of two seasons (2023 and 2024) make up the data. Duncan’s multiple range test indicates a significant difference at p ≤ 0.05 between the mean values ± SE of each parameter followed by various alphabetical letters. FC, field capacity; SE, standard error; TH, trehalose; V.CO, vermicompost.
The influence of diverse levels of drought (70% FC and 40% FC), V.CO and exogenous TH on chlorophyll a (μg · cm2 DW) (A), chlorophyll b (μg · cm2 DW) (B) and carotenoids (μg · cm2 DW) (C). Three replicates (n = 3) for each season and the means of two seasons (2023 and 2024) make up the data. Duncan’s multiple range test indicates a significant difference at p ≤ 0.05 between the mean values ± SE of each parameter followed by various alphabetical letters. FC, field capacity; SE, standard error; TH, trehalose; V.CO, vermicompost.

Figure 6.

The influence of diverse levels of drought (70% FC and 40% FC), V.CO and exogenous TH on Proline (mg · g−1 FW) (A), DPPH (%) (B), total phenolics (mg · g−1 DW) (C) and total flavonoids (mg · g−1 DW) (D). Three replicates (n = 3) for each season and the means of two seasons (2023 and 2024) make up the data. Duncan’s multiple range test indicates a significant difference at p ≤ 0.05 between the mean values ± SE of each parameter followed by various alphabetical letters. DPPH, 2,2-diphenyl-1-picrylhydrazyl; DW, dry weight; FC, field capacity; FW, fresh weight; SE, standard error; TH, trehalose; V.CO, vermicompost.
The influence of diverse levels of drought (70% FC and 40% FC), V.CO and exogenous TH on Proline (mg · g−1 FW) (A), DPPH (%) (B), total phenolics (mg · g−1 DW) (C) and total flavonoids (mg · g−1 DW) (D). Three replicates (n = 3) for each season and the means of two seasons (2023 and 2024) make up the data. Duncan’s multiple range test indicates a significant difference at p ≤ 0.05 between the mean values ± SE of each parameter followed by various alphabetical letters. DPPH, 2,2-diphenyl-1-picrylhydrazyl; DW, dry weight; FC, field capacity; FW, fresh weight; SE, standard error; TH, trehalose; V.CO, vermicompost.

Figure 7.

The influence of diverse levels of drought (70% FC and 40% FC), V.CO and exogenous TH on superoxide anion (O2) (mmole · min−1 · g−1 FW), (A), hydrogen peroxide (H2O2) (mmole · min−1 · g−1 FW) (B), IL (%) (C) and MDA (μmole · g−1 FW) (D). Three replicates (n = 3) for each season and the means of two seasons (2023 and 2024) make up the data. Duncan’s multiple range test indicates a significant difference at p ≤ 0.05 between the mean values ± SE of each parameter followed by various alphabetical letters. FC, field capacity; FW, fresh weight; IL, ion leakage; MDA, malondialdehyde; SE, standard error; TH, trehalose; V.CO, vermicompost.
The influence of diverse levels of drought (70% FC and 40% FC), V.CO and exogenous TH on superoxide anion (O2) (mmole · min−1 · g−1 FW), (A), hydrogen peroxide (H2O2) (mmole · min−1 · g−1 FW) (B), IL (%) (C) and MDA (μmole · g−1 FW) (D). Three replicates (n = 3) for each season and the means of two seasons (2023 and 2024) make up the data. Duncan’s multiple range test indicates a significant difference at p ≤ 0.05 between the mean values ± SE of each parameter followed by various alphabetical letters. FC, field capacity; FW, fresh weight; IL, ion leakage; MDA, malondialdehyde; SE, standard error; TH, trehalose; V.CO, vermicompost.

Figure 8.

The influence of diverse levels of drought (70% FC and 40% FC), V.CO and exogenous TH on CAT (mmole · min−1 · g−1 FW) (A) and APX (mmole · min−1 · g−1 FW) (B). Three replicates (n = 3) for each season and the means of two seasons (2023 and 2024) make up the data. Duncan’s multiple range test indicates a significant difference at p ≤ 0.05 between the mean values ± SE of each parameter followed by various alphabetical letters. APX, ascorbate peroxides; CAT, catalase; FC, field capacity; FW, fresh weight; SE, standard error; TH, trehalose; V.CO, vermicompost.
The influence of diverse levels of drought (70% FC and 40% FC), V.CO and exogenous TH on CAT (mmole · min−1 · g−1 FW) (A) and APX (mmole · min−1 · g−1 FW) (B). Three replicates (n = 3) for each season and the means of two seasons (2023 and 2024) make up the data. Duncan’s multiple range test indicates a significant difference at p ≤ 0.05 between the mean values ± SE of each parameter followed by various alphabetical letters. APX, ascorbate peroxides; CAT, catalase; FC, field capacity; FW, fresh weight; SE, standard error; TH, trehalose; V.CO, vermicompost.

Figure 9.

The transverse section through the leaves blade on the median portion of Volkamer lemon rootstock as affected by drought stress and anti stress (V.CO and TH) shows asymmetric. (A) Control with 100% FC; (B) with 70% FC; (C) with 40% FC; (D) with 70% FC + V.CO; (E) 40% F C + V.CO; (F) 70% FC + TH; (G) 40% FC + TH; (H) 70% FC + V.CO + TH; (I) 40% FC + V.CO + TH. (Obj. 10×). FC, field capacity; LE, lower epidermis; PT, palisade tissue; SD, schizogenous duct; ST, spongy tissue; TH, trehalose; UE, upper epidermis; VB, vascular bundle; V.CO, vermicompost.
The transverse section through the leaves blade on the median portion of Volkamer lemon rootstock as affected by drought stress and anti stress (V.CO and TH) shows asymmetric. (A) Control with 100% FC; (B) with 70% FC; (C) with 40% FC; (D) with 70% FC + V.CO; (E) 40% F C + V.CO; (F) 70% FC + TH; (G) 40% FC + TH; (H) 70% FC + V.CO + TH; (I) 40% FC + V.CO + TH. (Obj. 10×). FC, field capacity; LE, lower epidermis; PT, palisade tissue; SD, schizogenous duct; ST, spongy tissue; TH, trehalose; UE, upper epidermis; VB, vascular bundle; V.CO, vermicompost.

Figure 10.

The transverse section through the leaves blade on the median portion of Sour orange rootstock as affected by drought stress and anti stress (V.CO and TH) shows asymmetric. (A) Control with 100% FC; (B) with 70% FC; (C) with 40% FC; (D) with 70% FC + V.CO; (E) 40% FC + V.CO; (F) 70% FC + TH; (G) 40% FC + TH; (H) 70% FC + V. CO + TH; (I) 40% FC + V.CO + TH. (Obj. 10×). FC, field capacity; LE, lower epidermis; PT, palisade tissue; SD, schizogenous duct; ST, spongy tissue; TH, trehalose; UE, upper epidermis; V.CO, vermicompost; VB, vascular bundle.
The transverse section through the leaves blade on the median portion of Sour orange rootstock as affected by drought stress and anti stress (V.CO and TH) shows asymmetric. (A) Control with 100% FC; (B) with 70% FC; (C) with 40% FC; (D) with 70% FC + V.CO; (E) 40% FC + V.CO; (F) 70% FC + TH; (G) 40% FC + TH; (H) 70% FC + V. CO + TH; (I) 40% FC + V.CO + TH. (Obj. 10×). FC, field capacity; LE, lower epidermis; PT, palisade tissue; SD, schizogenous duct; ST, spongy tissue; TH, trehalose; UE, upper epidermis; V.CO, vermicompost; VB, vascular bundle.

Figure 11.

Leaf transverse section of Volkamer lemon rootstock as affected by drought stress and anti-stress (V.CO and TH) showing two unequal palisade parenchyma. (A) Control with 100% FC; (B) with 70% FC; (C) with 40% FC; (D) with 70% FC + V.CO; (E) 40% FC + V.CO; (F) 70% FC + TH; (G) 40% FC + TH; (H) 70% FC + V.CO + TH; (I) 40% FC + V.CO + TH (Obj. 10× and 40×). FC, field capacity; LE, lower epidermis; PT, palisade tissue; ST, spongy tissue; TH, trehalose; UE, upper epidermis; VB, vascular bundle; V.CO, vermicompost.
Leaf transverse section of Volkamer lemon rootstock as affected by drought stress and anti-stress (V.CO and TH) showing two unequal palisade parenchyma. (A) Control with 100% FC; (B) with 70% FC; (C) with 40% FC; (D) with 70% FC + V.CO; (E) 40% FC + V.CO; (F) 70% FC + TH; (G) 40% FC + TH; (H) 70% FC + V.CO + TH; (I) 40% FC + V.CO + TH (Obj. 10× and 40×). FC, field capacity; LE, lower epidermis; PT, palisade tissue; ST, spongy tissue; TH, trehalose; UE, upper epidermis; VB, vascular bundle; V.CO, vermicompost.

Figure 12.

Leaf transverse section of sour orange rootstock as affected by drought stress and anti-stress (V.CO and TH) showing two unequal palisade parenchyma. (A) Control with 100% FC; (B) with 70% FC; (C) with 40% FC; (D) with 70% FC + V.CO; (E) 40% FC + V.CO; (F) 70% FC + TH; (G) 40% FC + TH; (H) 70% FC + V.CO + TH; (I) 40% FC + V.CO + TH (Obj. 10× and 40×). FC, field capacity; LE, lower epidermis; PT, palisade tissue; SD, schizogenous duct; ST, spongy tissue; TH, trehalose; UE, upper epidermis; VB, vascular bundle; V.CO, vermicompost.
Leaf transverse section of sour orange rootstock as affected by drought stress and anti-stress (V.CO and TH) showing two unequal palisade parenchyma. (A) Control with 100% FC; (B) with 70% FC; (C) with 40% FC; (D) with 70% FC + V.CO; (E) 40% FC + V.CO; (F) 70% FC + TH; (G) 40% FC + TH; (H) 70% FC + V.CO + TH; (I) 40% FC + V.CO + TH (Obj. 10× and 40×). FC, field capacity; LE, lower epidermis; PT, palisade tissue; SD, schizogenous duct; ST, spongy tissue; TH, trehalose; UE, upper epidermis; VB, vascular bundle; V.CO, vermicompost.

Lamina anatomical traits of citrus rootstock leaves as affected by drought stress and anti-stress (V_CO and TH)_

Rootstock typeTreatment no.Cuticle thickness (μm)Lamina thickness (μm)Midrib zone thickness (μm)Cell size (μm)VB of mid rib (μm)
Palisade mesophyllSpongy mesophyllVertical lengthHorizontal length
Volkamer lemonT197.39 ± 0.008 c15.44 ± 5.773 a61.17 ± 5.773 c3.41 ± 5.773 d10.97 ± 3.333 d39.79 ± 5.773 a45.57 ± 3.333 c
T267.02 ± 0.014 h12.84 ± 0.010 h54.86 ± 6.666 f2.36 ± 5.773 i9.62 ± 3.333 h34.32 ± 5.773 g46.31 ± 5.773 a
T370.87 ± 0.014 g12.18 ± 0.003 i51.57 ± 5.773 h3.11 ± 5.773 g10.94 ± 6.666 e34.35 ± 8.819 f43.12 ± 8.819 g
T466.37 ± 0.037 i13.03 ± 6.666 g56.82 ± 8.819 d2.54 ± 8.190 h9.57 ± 6.666 i34.02 ± 5.773 h39.11 ± 3.333 h
T580.38 ± 0.008 f14.13 ± 3.333 f54.16 ± 0.002 g4.58 ± 5.773 a11.09 ±6.666 a36.06 ±0.001 e44.66 ± 44.66 d
T684.09 ±0.011 e14.27 ± 5.773 e55.55 ± 6.666 e3.62 ± 5.773 b10.99 ± 6.666 b36.49 ± 5.773 d43.35 ± 43.35 f
T788.81 ±0.017 d14.30 ± 8.819 d50.09 ± 6.666 i3.14 ± 8.819 f10.98 ± 3.333 c30.97 ± 5.773 i37.45 ± 5.773 i
T8100.00 ±0.021 b14.88 ±0.001 c61.98 ± 3.333 a3.46 ± 6.666 c10.66 ± 0.003 g37.20 ± 5.773 c45.88 ± 3.333 b
T9102.16 ±0.005 a15.01 ± 8.819 b61.71 ± 8.819 b3.32 ± 8.819 e10.66 ± 0.001 f38.61 ± 5.773 b44.38 ±0.001 e
Sour orangeT190.12 ± 0.014 c18.02 ± 5.773 a57.29 ± 5.773 a3.77 ± 0.148 c13.88 ±5.773 a36.75 ± 0.001 c43.90 ± 8.819 e
T272.14 ± 0.012 h13.84 ± 8.819 h46.30 ± 5.773 h2.71 ± 3.333 e10.97 ± 8.819 h29.39 ± 5.773 g37.16 ± 3.333 g
T369.58 ±0.011 i12.90 ± 8.819 i42.69 ± 0.003 i3.29 ± 5.773 d8.69 ± 5.773 i25.15 ± 5.773 h33.58 ± 3.333 h
T475.25 ± 0.017 g16.69 ± 3.333 e55.70 ± 5.773 e3.82 ± 5.773 b11.41 ± 5.773 g35.68 ± 5.773 d46.23 ± 8.819 c
T580.31 ± 0.008 f16.82 ± 5.773 d43.32 ± 0.003 g2.85 ± 8.819 e12.02 ±0.001 e24.13 ± 5.773 i32.41 ± 3.333 i
T683.47 ± 0.014 e15.32 ± 5.773 g59.23 ± 8.819 c3.59 ±0.001 c11.90 ± 5.773 f35.45 ± 5.773 e41.82 ± 8.819 f
T787.35 ± 0.014 e15.60 ± 8.819 f52.84 ± 8.189 f3.89 ± 3.333 b12.90 ±0.001 c33.46 ± 5.773 f44.07 ± 3.333 d
T894.82 ± 0.014 b17.09 ± 6.666 c60.87 ± 5.773 b4.09 ± 5.773 a12.34 ± 5.773 d41.36 ±0.001 b51.23 ± 5.773 b
T997.32 ±0.014 a17.77 ± 5.773 b72.42 ± 5.773 a3.74 ± 3.333 be13.27 ± 5.773 b45.36 ±0.001 a64.20 ± 3.333 a

Lamina micro-morphological characters of citrus rootstock leaves as affected by drought stress and anti-stress (V_CO and TH)_

Dermal systemMesophyll tissueMechanical tissueGround tissueVascular systemSecretory systemCrystals
Trichomes
Rootstock typeTreatment no.Outline in T.sEglandularGlandularCuticleEpidermal cellsTypePalisade Rows No.Palisade extended at mid rib regionCollenchymaParenchymaOutlineNo of bundlesSchizogenous duct
Volkamer lemonTlRaised adaxially & abaxially--ThickTangentially, radiallyDorsiventral2+Annular+Continuous37+-
T2//--Thin//////+//+//45+-
T3//--////////+//+//34+-
T4//--////////+//+//26+-
T5//--Thick//////+//+//25+-
T6//--////////+//+//23+-
T7//--////////+//+//23+-
T8//--////////+//+//24+-
T9//--////////+//+//36+-
Sour orangeTl//--Thick//////+//+//40+-
T2//--Thin//////+//+//52+-
T3//--////////+//+//46+-
T4//--Thick//////+//+//58+-
T5//--////////+//+//37+-
T6//--////////+//+//35+-
T7//--////////+//+//37+-
T8//--////////+//+//39+-
T9//--////////+//+//38+-
DOI: https://doi.org/10.2478/fhort-2025-0023 | Journal eISSN: 2083-5965 | Journal ISSN: 0867-1761
Language: English
Submitted on: Apr 7, 2025
Accepted on: Nov 1, 2025
Published on: Dec 12, 2025
Published by: Polish Society for Horticultural Sciences (PSHS)
In partnership with: Paradigm Publishing Services
Publication frequency: 2 issues per year

© 2025 M. S. Aboryia, Amr Elkelish, Lina M. Abu-Ziada, Mai M. Wahba, Abdulrahman Alhudhaibi, Mohamed S. Gawish, published by Polish Society for Horticultural Sciences (PSHS)
This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivatives 3.0 License.

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