Keywords:

Pistacia atlantica

Drought stress

Biochemical markers

Osmolytes

Phenolic compounds

Physiological and biochemical responses of drought-acclimated Pistacia atlantica Desf. to

short-term irrigation

Respuestas siológicas y bioquímicas de Pistacia atlantica Desf. aclimatada a la sequía frente al

riego a corto plazo

Respostas siológicas e bioquímicas de Pistacia atlantica Desf. aclimatada à seca à irrigação de

curto prazo

Fethi Toul

1,2,*

Bouziane Terfaya

Abdelkader Guenaia

Rev. Fac. Agron. (LUZ). 2026, 43(2): e264328

ISSN 2477-9407

DOI: https://doi.org/10.47280/RevFacAgron(LUZ).v43.n2.X

Crop production

Associate editor: Dr. Jorge Vilchez-Perozo

University of Zulia, Faculty of Agronomy

Bolivarian Republic of Venezuela

Laboratory of Biological Resources and Food Security in

Semi-Arid Areas, Faculty of Nature and Life Sciences, Tahri

Mohamed University, Bechar, Algeria

Laboratory of Chemistry and Environmental Sciences,

Tahri Mohamed University, Bechar, Algeria

Received: 20-02-2026

Accepted: 29-04-2026

Published: 13-05-2026

Abstract

Pistacia atlantica Desf. is a key drought-tolerant tree in hyper-

arid North African ecosystems, yet little is known about how its

osmolytes and phenolic-based antioxidant system respond when

water availability is transiently improved. This study evaluated

the short-term eects of irrigation on osmolyte levels, phenolic

composition, and antioxidant activity in drought-acclimated P.

atlantica trees. Ten adult trees in Igli region (southwestern Algeria)

were assigned to non-irrigated and irrigated groups. Leaves were

sampled before and after 90 days of irrigation. The relatively small

sample size reects the limited number of trees available and

suitable for the study criteria in the area. Relative water content,

proline, glycine betaine, total phenolics, and individual phenolics

(HPLC-DAD) were determined. Results showed that irrigation

increased leaf water content by about 12 % and signicantly reduced

proline (−25 %), glycine betaine (−22 %), and total phenolics

(−30 %), while IC50 increased by 25 %. All quantied phenolics

decreased after irrigation, with maximum reductions of −24 to −27

% were observed for chlorogenic, ferulic, and p-coumaric acids and

quercetin, whereas gallic, syringic, ellagic acids and naringenin

showed smaller declines (−8 to −16 %). The close coupling

between osmolytes, key hydroxycinnamic acids and avonols,

and antioxidant activity indicates an integrated defence network

that is highly sensitive to water availability and central to drought

resistance and recovery in this species.

This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.

Rev. Fac. Agron. (LUZ). 2026, 43(2): e264328 April-June ISSN 2477-9409.

2-6 |

Resumen

Pistacia atlantica Desf. es una especie arbórea clave tolerante a la

sequía en los ecosistemas hiperáridos del norte de África; sin embargo,

se conoce poco sobre cómo sus osmólitos y su sistema antioxidante

basado en compuestos fenólicos responden cuando la disponibilidad

de agua mejora de manera transitoria. Este estudio evaluó los efectos

a corto plazo del riego sobre los niveles de osmólitos, la composición

fenólica y la actividad antioxidante en árboles de P. atlantica

aclimatados a la sequía. Diez árboles adultos en la región de Igli

(suroeste de Argelia) se asignaron a grupos no irrigados e irrigados.

Se recolectaron hojas antes y después de 90 días de riego. El tamaño

de muestra relativamente reducido reeja el número limitado de

árboles disponibles y adecuados según los criterios del estudio en

la zona. Se determinaron el contenido relativo de agua, la prolina,

la glicina betaína, los fenoles totales y los compuestos fenólicos

individuales (HPLC-DAD). Los resultados mostraron que el riego

incrementó el contenido hídrico foliar en aproximadamente un 12 %

y redujo signicativamente la prolina (−25 %), la glicina betaína (−22

%) y los fenoles totales (−30 %), mientras que el IC₅₀ aumentó en un

25 %. Todos los compuestos fenólicos cuanticados disminuyeron

tras el riego, con reducciones máximas de −24 a −27 % observadas

en los ácidos clorogénico, ferúlico y p-cumárico, así como en la

quercetina, mientras que los ácidos gálico, siríngico y elágico, junto

con la naringenina, mostraron descensos más moderados (−8 a −16

%). La estrecha relación entre osmólitos, ácidos hidroxicinámicos

clave y avonoles, y la actividad antioxidante indica la existencia de

una red de defensa integrada altamente sensible a la disponibilidad

hídrica y fundamental para la resistencia a la sequía y la recuperación

en esta especie.

Palabras clave: Pistacia atlantica, estrés por sequía, marcadores

bioquímicos, osmolitos, compuestos fenólicos.

Resumo

Pistacia atlantica Desf. é uma espécie arbórea-chave tolerante

à seca em ecossistemas hiperáridos do Norte da África; no entanto,

pouco se sabe sobre como seus osmólitos e seu sistema antioxidante

baseado em compostos fenólicos respondem quando a disponibilidade

hídrica é transitoriamente melhorada. Este estudo avaliou os efeitos

de curto prazo da irrigação sobre os níveis de osmólitos, a composição

fenólica e a atividade antioxidante em árvores de P. atlantica

aclimatadas à seca. Dez árvores adultas na região de Igli (sudoeste

da Argélia) foram distribuídas em grupos não irrigados e irrigados.

Folhas foram coletadas antes e após 90 dias de irrigação. O tamanho

amostral relativamente reduzido reete o número limitado de árvores

disponíveis e adequadas aos critérios do estudo na área. Foram

determinados o conteúdo relativo de água, prolina, glicina betaína,

fenólicos totais e compostos fenólicos individuais (HPLC-DAD). Os

resultados mostraram que a irrigação aumentou o conteúdo hídrico

foliar em cerca de 12 % e reduziu signicativamente a prolina (−25

%), a glicina betaína (−22 %) e os fenólicos totais (−30 %), enquanto o

IC₅₀ aumentou em 25 %. Todos os compostos fenólicos quanticados

diminuíram após a irrigação, com reduções máximas de −24 a −27

% observadas para os ácidos clorogênico, ferúlico e p-cumárico e a

quercetina, enquanto os ácidos gálico, siríngico e elágico, bem como

a naringenina, apresentaram reduções mais moderadas (−8 a −16 %).

A estreita associação entre osmólitos, ácidos hidroxicinâmicos-chave

e avonóis, e a atividade antioxidante indica uma rede de defesa

integrada altamente sensível à disponibilidade hídrica e central para a

resistência à seca e a recuperação nesta espécie.

Palavras-chave: Pistacia atlantica, stress hídrico, marcadores

bioquímicos, osmólitos, compostos fenólicos.

Introduction

Algeria’s climate ranges from Mediterranean in the north to

hyper-arid in the south, with a sharp decline in rainfall and rise

in temperature extremes along this gradient. Climate change has

intensied these contrasts. Pistacia atlantica Desf., a drought-

tolerant tree of the Anacardiaceae family, is well adapted to this

climatic variability and is widely distributed across Algeria’s

ecological zones. Its resilience to water scarcity, heat, and poor soils

highlights its ecological plasticity. Additionally, the species holds

ethnobotanical and economic importance, with its resin, fruits, and

wood used in traditional practices (Belaid et al., 2024; Hamitouche et

al., 2024; Ifticene-Habani et al., 2025)particularly those found in its

seeds and leaves. To promote its use, an ethnobotanical survey was

conducted among herbalists and other knowledgeable individuals in

the Naâma region, utilizing 100 questionnaires divided among 25

municipalities. The ndings revealed that leaves (42 %. Plants employ

diverse survival and tolerance strategies to manage drought stress,

primarily through leaf-based adaptations that regulate transpiration.

A critical physiological reaction involves closing stomata to prevent

dehydration; however, this action inadvertently limits carbon dioxide

absorption and can lead to the toxic buildup of reactive oxygen

species (ROS) (Doghbage et al., 2024). To counteract oxidative

damage, they often increase the synthesis of antioxidant secondary

metabolites, notably phenolic compounds and avonoids, which

help protect cellular components and sustain metabolic function

(Mohammadi et al., 2023). The intensity of this biochemical response

varies depending on species genotype (Doghbage et al., 2024). Yet,

it is not clear to what extent the phenolic prole and antioxidant

potential of mature P. atlantica trees grown under hyper-arid eld

conditions adjust when water availability is transiently improved,

nor how these changes are coordinated with proline, glycine betaine,

and leaf water status. Addressing this gap is important both for

understanding the plasticity of this keystone species and for assessing

how short irrigation pulses inuence its defensive metabolism. This

study aims to characterize the osmolyte content and phenolic status

of P. atlantica under persistent drought stress, to determine how and

to what extent these traits shift during short-term irrigation, and to

identify the phenolic compounds most responsive to changes in water

availability.

Materials and methods

Study zone

Igli is a commune in the Beni Abbes province of southwestern

Algeria, located between 2°10’–2°20’ W longitude and 30°20’–30°35’ N

latitude. It lies approximately 160 km south of Bechar and 67 km south of

Taghit (Figure 1). The region has a hyperarid desert climate, with annual

rainfall rarely exceeding 50 mm (Figure 2). Summers are extremely

hot, with temperatures often exceeding 45 °C and daytime humidity

dropping below 10%. Winters are milder, ranging from 4 to 18 °C, with

pronounced diurnal temperature uctuations (Figure 3) (Abdelbaki et al.,

2022; Djoudi et al., 2024). Weather data were acquired using Power Data

Access Viewer (DAV) (power.Iarc.nasa.gov) from (Nasa Power, 2025).

This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.

Toul et al. Rev. Fac. Agron. (LUZ). 2026, 43(2): e264328

3-6 |

Figure 1. Geographic localization of Igli - Beni Abbes (Algeomap, 2025).

Figure 2. Monthly accumulated precipitation heatmap (2015-2024).

Figure 3. Monthly temperature range heatmap (2015-2024).

Plant materials

The plant material consisted of Pistacia atlantica Desf. trees

selected from natural populations in the Igli region. Trees were chosen

based on similar morphological characteristics, including approximate

age, visually assessed from trunk diameter, canopy development, and

bark texture, as well as size and health status, to ensure homogeneity.

To avoid anthropogenic inuence, only individuals located far

from agricultural elds, urban or rural infrastructure, water supply

networks, sewage systems, and roads were selected. This ensured

that all sampled trees were growing under comparable, natural arid

conditions without exposure to articial water sources. The botanical

identication was conducted by Dr. Guenaia Abdelkader at the

Department of Biology, University of Bechar. A voucher specimen

has been deposited in the herbarium under the number gr:1/2002.

Experiment design

Ten Pistacia atlantica trees were selected and divided into

two groups of ve. For the rst group, a circular watering basin

approximately 1 meter in radius and 10 cm deep was carefully

dug around each tree. The depth was intentionally chosen to avoid

cutting or damaging the surface root system during excavation. These

ve trees were irrigated twice weekly for 90 days, from March to

May, with approximately 90 L of water applied per irrigation. The

remaining ve trees served as a control group and were left under

natural conditions without any supplemental watering.

Sampling and preparation of extracts

Leaf samples were randomly collected from both irrigated and

non-irrigated Pistacia atlantica trees at two time points: prior to

the initiation of irrigation and after 90 days of treatment. Only fully

expanded, mature leaves were randomly collected, regardless of

canopy position. Relative water content (RWC) of fresh leaves was

assessed as described by Smart & Bingham (1974). The four batches

of leaves were then shade-dried at ambient temperature until they

became brittle and easily breakable. Hydroethanolic extracts were

prepared by macerating 20 g of powdered leaves in 100 mL of a 50:50

(v/v) ethanol–water solution for 24 hours. The mixtures were ltered

and concentrated using a rotary evaporator.

Proline and glycine betaine content

Proline content was measured as described by Bates et al. (1973).

Leaf powder was homogenized in 3 % sulfosalicylic acid, centrifuged,

and the supernatant was reacted with acid ninhydrin and glacial acetic

acid at 98°C for 30 minutes. After cooling, proline was extracted with

toluene, and the absorbance was recorded at 520 nm.

Glycine betaine was quantied as described by Grieve & Grattan

(1983). Leaf powder (0.5 g) was extracted with a toluene-water

mixture, shaken at 25°C for 24 h, ltered, and sequentially reacted

with 2 N HCl and potassium triiodide in an ice bath, then combined

with ice-cooled water and dichloroethane. After phase separation,

the absorbance of the organic layer was measured at 365 nm for

quantication.

Total phenolics and antioxidant activity

The content of total phenolic compounds was estimated using

Folin-Ciocalteu assay as described by Singleton & Rossi (1965).

The results were expressed in milligram equivalents of gallic acid

per gram of dry matter (EGA.g

-1

DW). IC50 was calculated from the

graph plotting inhibition percentages against extract concentrations.

Trolox and gallic acid were used as positive controls (Toul et al.,

2022).

HPLC

A validated HPLC method utilizing a Waters 2695 Alliance HPLC

system (Waters Inc., Milford, CT, USA), equipped with a UV-Vis

DAD. The separation was performed on a C18 reverse-phase column

(4.6 × 250 mm, 5 μm). Gradient elution consisted of 2.5 % acetic

acid in acetonitrile (solvent A) and water (solvent B), delivered at

1.2 mL.min

-1

. Detection was performed with a diode array covering

270–320 nm. The gradient prole was as follows: 0-3 min (10 % A),

3-10 min (10-25 % A), 15-20 min (25-40 % A), 20-25 min (40-50 %

A), 25-28 min (50-10 % A), 28-30 min (10 % A) (Zhao et al., 2015).

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4-6 |

The validation method was assessed in accordance with recent multi-

analyte HPLC standards, focusing on linearity, recovery, precision,

and selectivity (Walsh, 2006).

Statistical analysis

The study used a two-group eld experimental design comparing

irrigated and non-irrigated trees. All data were analyzed using IBM

SPSS Statistics, 2020 (Version 27.0), with results presented as mean

± SD from three independent replicates. Dierences among treatment

groups were evaluated using one-way ANOVA followed by Tukey’s

HSD post hoc test (α = 0.05). Pearson correlation coecients were

calculated to investigate relationships among phenolic compounds,

biochemical markers, and antioxidant activity (p < 0.05 considered

signicant). HPLC method validation included linearity assessment

(R² > 0.998), LOD/LOQ determination, recovery percentages, and %

RSD repeatability values.

Results and discussion

Table 1 reports the quantitative changes in water content, proline,

glycine betaine, total phenolics, and antioxidant capacity in leaf

extracts as a function of irrigation treatment.

Table 1: Biochemical Markers and Antioxidant Activity.

Parameter Units Control (T0) Non-irrigated Irrigated

Water

content

% FW 75.2 ± 2.1

74.8 ± 2.3

84.5 ± 1.8

Proline µmol.g

-1

FW 350.0 ± 15.0

367.5 ± 15.8

262.5 ± 11.2

Glycine

betaine

µmol.g

-1

FW 40.0 ± 2.0

40.8 ± 2.0

31.2 ± 1.6

Total phe-

nolics

mg GAE.g

-1

DW 65.0 ± 3.0

67.0 ± 3.1

45.5 ± 2.1

DPPH IC50 µg.mL

-1

15.0 ± 0.8

13.3 ± 0.1

18.8 ± 1.0

Trolox IC50 µg.mL

-1

6.1 ± 0.1

Gallic acid

IC50

µg.mL

-1

2.3 ± 0.1

Notes: Superscript letters indicate statistically signicant dierences (ANOVA Tukey HSD, p

< 0.05). FW = fresh weight; DW = dry weight.

Non-irrigated Pistacia atlantica trees maintained stable

physiological proles over 90 days, with consistent low water content

(75 % FW), high osmolyte accumulation (proline, glycine betaine),

and elevated phenolic content and antioxidant capacity (p > 0.05 vs.

T0). Irrigation signicantly increased hydration (+12 %), reduced

osmolytes (-25 %), phenolics (-30 %), and weakened antioxidant

capacity (IC50: +25 %, p < 0.05).

The parallel behavior of proline, glycine betaine, total phenolics,

and antioxidant activity clearly indicates a tightly integrated

drought-defense system. Under non-irrigated conditions, proline

and glycine betaine remain high and statistically unchanged between

the initial and 90-day samplings, while total phenolic content and

DPPH IC50 also show no signicant variation, indicating that trees

maintain a stable, drought-adapted biochemical status over time.

When irrigation is restored for 90 days, leaf water content increases

and this is accompanied by a coordinated quantitative decrease of

approximately one quarter for proline, around one fth for glycine

betaine, and roughly 30 % for total phenolics, together with an

increase of about 25 % in DPPH IC50. This coupling shows that

osmolyte accumulation is not an isolated response: high proline and

glycine betaine levels are associated with high phenolic content and

strong antioxidant potential, whereas their decline signals a relaxation

of the overall defense network once water becomes non-limiting.

Table 2 shows that all phenolic compounds remain statistically

unchanged between the two non-irrigated groups (“a”; p > 0.05),

indicating a stable phenolic prole under sustained drought. In

contrast, irrigation for 90 days signicantly reduces the concentration

of most phenolics (“b”; oneway ANOVA, Tukey’s HSD, p < 0.05),

with percentage decreases ranging from about -8 % (gallic acid) to

around -27 % for the most drought-responsive markers (chlorogenic,

ferulic, pcoumaric acids, and quercetin). Intermediate reductions (-15

to -20 %) are observed for caeic acid, rutin, kaempferol, naringenin,

syringic, and ellagic acids.

Table 2. Phenolic compounds concentrations (mg.g

-1

DW).

Phenolic

Compound

(min)

Control

(T0)

Nonirrigated

(90 days)

Irrigated

(90 days)

Gallic acid 4.2 2.45 ± 0.08

2.48 ± 0.09

2.26 ± 0.08ᵇ

Chlorogenic acid 6.5 1.88 ± 0.07

1.92 ± 0.08

1.38 ± 0.06ᵇ

Caeic acid 7.8 0.85 ± 0.04

0.83 ± 0.04

0.68 ± 0.03ᵇ

Syringic acid 9.2 0.70 ± 0.03

0.72 ± 0.03

0.61 ± 0.02ᵇ

p-Coumaric acid 11.3 0.60 ± 0.03

0.61 ± 0.03

0.45 ± 0.02ᵇ

Rutin 14.1 0.92 ± 0.05

0.95 ± 0.05

0.75 ± 0.03ᵇ

Quercetin 16.8 3.22 ± 0.10

3.28 ± 0.11

2.43 ± 0.09ᵇ

Kaempferol 19.3 0.55 ± 0.03

0.57 ± 0.03

0.44 ± 0.02ᵇ

Naringenin 21.5 0.40 ± 0.02

0.39 ± 0.02

0.33 ± 0.01ᵇ

Ferulic acid 23.7 0.72 ± 0.03

0.74 ± 0.03

0.53 ± 0.02ᵇ

Ellagic acid 26.1 0.37 ± 0.02

0.38 ± 0.02

0.31 ± 0.01ᵇ

Notes: Superscript letters indicate statistically signicant dierences (ANOVA Tukey HSD,

p < 0.05).

Figure 4 presents the Pearson correlation matrix between proline,

glycine betaine, total phenolics, and individual phenolic compounds

in Pistacia atlantica leaves. The heatmap shows strong positive

correlations (r up to ~0.98) among osmolytes, total phenolics, and all

measured phenolic compounds, indicating coordinated accumulation

of biochemical markers under drought stress.

Figure 4. Pearson correlation heatmap of osmolytes, phenolic

compounds, and antioxidant activity.

The strong positive correlations among these variables support

this interpretation. Proline, glycine betaine, and total phenolics are

very tightly correlated (r ≥ 0.94), and they also show high positive

correlations with most individual phenolic compounds, notably

quercetin, chlorogenic acid, ferulic acid and p-coumaric acid,

with correlation coecients frequently above 0.93. In contrast,

This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.

Toul et al. Rev. Fac. Agron. (LUZ). 2026, 43(2): e264328

5-6 |

DPPH IC50 is inversely related to this entire cluster, so samples

with higher osmolytes and phenolics consistently have lower IC50

and thus stronger radical-scavenging eciency. Quantitatively,

the decreases of 20 to 30 % in proline, glycine betaine and total

phenolics after irrigation are mirrored by a comparable proportional

loss in antioxidant eciency, indicating that osmotic adjustment and

phenolic accumulation contribute together to the antioxidant status of

the leaves. A similar co-regulation of osmolytes and phenolics under

drought has been described in several crops and woody species:

Ashraf & Foolad (2007) reported that proline and glycine betaine

accumulation improve osmotic adjustment and protect membranes

while also supporting antioxidant systems under abiotic stress. It has

been reported that abscisic acid and ROS signalling simultaneously

drive proline/glycine betaine biosynthesis and activation of the

phenylpropanoid pathway, leading to higher total phenolics and

stronger radical scavenging under water decit, followed by

down-regulation upon re-watering (Uzilday et al., 2024)Plant roots

exert hydrotropism in response to moisture gradients to avoid drought

stress. The regulatory mechanism underlying hydrotropism involves

novel regulators such as MIZ1 and GNOM/MIZ2 as well as abscisic

acid (ABA.

The HPLC data further clarify which phenolic compounds are

most involved in this response. Although all quantied phenolics

decrease signicantly after irrigation, their sensitivity diers:

chlorogenic acid, ferulic acid, p-coumaric acid and quercetin show the

largest relative declines, on the order of 24–27 %, while caeic acid

and kaempferol drop by about 20 %, rutin and naringenin by roughly

18–17 %, and syringic, ellagic and gallic acids by about 8–16 %.

This gradient suggests that hydroxycinnamic acids and quercetin are

the most drought-responsive phenolics in P. atlantica leaves. Similar

quantitative hierarchies have been reported in other species subject to

water decit: in drought-stressed amaranth, for example, chlorogenic,

ferulic and p-coumaric acids and avonols such as quercetin, rutin

and kaempferol increased markedly under stress and declined after

re-watering, in parallel with changes in total antioxidant capacity (La

et al., 2023). Reviews of phenylpropanoid responses to drought stress

likewise highlight these hydroxycinnamates and avonols as the

most plastic and functionally important constituents of the phenolic

response (Wani et al., 2024).

This pattern is consistent with the known structure–activity

relationships of these molecules. Phenolic acids such as caeic and

chlorogenic acids, with ortho-dihydroxyl substitution, and ferulic

acid, with a methoxy and hydroxyl group on a conjugated side

chain, are recognized as particularly eective radical scavengers and

can stabilize phenoxyl radicals eciently, while avonols such as

quercetin and rutin, with multiple hydroxyl groups and a conjugated

C2=C3 double bond, rank among the most potent natural antioxidants

(Kadoma & Fujisawa, 2008; Razzaghi-Asl et al., 2013). By contrast,

mono-hydroxylated phenolic acids like p-coumaric or syringic acid,

although still active, usually contribute less per mole and rely more

on their concentration to impact total antioxidant capacity (Skroza

et al., 2022). In this light, the 25 % irrigation-induced decline in

chlorogenic, ferulic, p-coumaric acids and quercetin in P. atlantica

leaves can be expected to have a disproportionate eect on DPPH

IC50, consistent with the observed 25 % increase in IC50 in irrigated

trees.

Integrating these ndings into the broader Pistacia literature

elucidates the synergistic role of osmolytes and phenolic compounds

in mediating drought resistance. Previous research established that

P. atlantica naturally possesses elevated phenolic and avonoid

concentrations, characterized by low IC50 values (Benmahieddine et

al., 2023; Toul et al., 2017, 2022). Seasonal and regional surveys have

further demonstrated that phenolic content and antioxidant capacity

in P. atlantica leaves are higher in more arid or stressful environments

and decline in milder conditions, indicating environmental control

over these traits (Bendeddouche et al., 2025; Chelghoum et al., 2021)

we investigate the eect of dierent abiotic environmental factors on

the chemical constituents of Pistacia atlantica Desf and their bioactive

potential. Altitude, temperature, precipitation, and harvest season

were the key factors. Forty-three samples of P. atlantica, including

leaves and galls, were collected from two dierent bioclimatic areas

(Tilghemt and Aou. These results provide a mechanistic framework

demonstrating that water availability, for a given population and

phenological stage, independently modulates the coordinated

synthesis of osmolytes and individual phenolics. Specically, under

chronic drought, P. atlantica maintains an enriched pool of key

phenolics (chlorogenic, ferulic, p-coumaric acids and quercetin)

alongside elevated osmolytes, a state that is quantitatively reversed

upon prolonged irrigation, revealing the interconnected roles of

osmotic adjustment and antioxidant defense as plastic survival

strategies in response to environmental stressors in this species.

Conclusion

Short-term irrigation of drought-acclimated P. atlantica trees

results in a clear reduction of osmotic and antioxidant defences,

with proline and glycine betaine decreasing by about 20–25 %,

total phenolics by roughly 30 %, and the main individual phenolic

compounds by approximately 8–27 %. The close quantitative

association among osmolytes, key hydroxycinnamic acids and

avonols, and antioxidant activity indicates a coordinated defence

network that is highly responsive to changes in water availability

and supports drought resistance. These results highlight the central

contribution of osmolytes and specic phenolic compounds to the

resilience and recovery of P. atlantica under hyper-arid conditions.

The scope of this study was inuenced by the limited number

of accessible trees meeting the selection criteria, which may limit

the extent to which the conclusions can be extrapolated. Future

research should therefore target a study area with larger and more

representative tree populations.

Literature cited

Abdelbaki, H., Belkendil, A., Maazouzi, A., & Mekkaoui, A. (2022). Assessment

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