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Article

Understanding the Wood Pocket Physiopathy in Persian Lime Through Its Physiological Characterization

by
Felipe Roberto Flores-de la Rosa
1,2,
Gabriela Fuentes-Ortíz
3,
Ricardo Santillán-Mendoza
1,
Cristian Matilde-Hernández
1,
Humberto Estrella-Maldonado
1,* and
Jorge M. Santamaría
2,*
1
Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias (INIFAP), CIR Golfo Centro, Campo Experimental Ixtacuaco, Km 4.5 Carretera Martínez de la Torre-Tlapacoyan, Tlapacoyan 93600, Veracruz, Mexico
2
Centro de Investigación Científica de Yucatán A.C., Calle 43 No. 130, Colonia Chuburná de Hidalgo, Mérida 97205, Yucatán, Mexico
3
Independant Researcher, Jardines de Vista Alegre, Mérida 97138, Yucatán, Mexico
*
Authors to whom correspondence should be addressed.
Agronomy 2025, 15(4), 762; https://doi.org/10.3390/agronomy15040762
Submission received: 24 January 2025 / Revised: 14 March 2025 / Accepted: 18 March 2025 / Published: 21 March 2025
(This article belongs to the Special Issue Physiological, Biochemical, and Molecular Mechanisms of Abiotic Stress Tolerance in Fruits)
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Abstract

:
Persian lime is one of the most important citrus fruits in Mexico. This species suffers from a physiopathy called Wood Pocket (WP), which is characterized by chlorosis in leaves and fruits, necrosis in the trunk, and the eventual death of the tree. The actual scientific knowledge about WP is very little; however, the producers, researchers, and technicians associate it with abiotic stress. In the present study, we observed that Persian limes exposed to temperatures over 45 °C for five months developed WP symptoms, while those kept at 25 °C during the same time did not. Both groups were then physiologically characterized. Our results show that WP strongly affects most fluorescence and gas exchange parameters. Specially, we observe that stomatal fluxes were negative in WP leaves. Our results suggest that WP in Persian lime plants might be a physiological affectation caused by the prolonged exposure to high temperatures. It might be possible that the physiological affectation (gas exchange and water use efficiency) could be associated with a high-temperature-induced abscisic acid accumulation, which, in turn, might have caused stomatal closure in WP-affected plants. However, this hypothesis must be corroborated experimentally.

1. Introduction

The Persian lime (Citrus latifolia Tanaka) is a highly consumed citrus fruit worldwide, often consumed fresh [1], but it also has many industrial uses, such as juice extraction and essential oil production [2]. This fruit tree is cultivated in tropical regions, with Mexico being the world’s leading producer and exporter [3]. Most of the producing fields in Mexico have low levels of technological advancement, relying mainly on seasonal conditions [4,5].
The ecological conditions predispose Persian lime to a variety of stress sources that directly impact productivity and production quality, such as biotic stress: fungal diseases [6], bacterial infections [7]; and abiotic stress: water stress [8], oleocellosis [9], among others. However, a condition of international interest has recently re-emerged, characterized by the appearance of chlorotic mottling on the leaves and sectoral spotting on the fruits [10], known as Wood Pocket (WP).
This condition was first reported in California in the 1940s [11], and in Florida in the 1950s [12], demonstrating that it was not the result of any microorganism infection but rather an intrinsic response of the plant to environmental conditions [13]. However, it is only in recent years that it has gained global relevance due to its impact on Persian lime production. Consequently, there are very few reports in the scientific literature regarding its characterization and nature. This disease has already been reported in Mexico [10,14], but basic knowledge about this physiopathy remains scarce. However, estimates performed by our group in the main Persian-lime-producing regions of Mexico indicate that the WP incidence in Persian-lime-producing areas exceeds 80% and varies in severity, with no observable pattern (unpublished data). Therefore, the present study focused on the physiological characterization of leaves with WP symptoms, laying the foundation for a better understanding of this physiopathy.

2. Materials and Methods

2.1. Plant Material

Eight-month-old Persian lime plants grafted onto C. volkameriana rootstock were used in this study. These plants were acquired from a certified nursery to prevent contamination by any phytopathogens according to NOM-079-FITO-2002 (https://www.gob.mx/senasica/documentos/nom-079-fito-2002 (accessed on 25 January 2024)). The plants were maintained from February to July 2024 under two different conditions: The first group was kept in a greenhouse with controlled temperature (25 ± 2 °C), with ambient CO2 levels ranging from 100 to 1500 ppm, and light intensity (photosynthetic photon flux density, PPFD) varying between 90 and 420 µmol PPFD m−2 s−1. On the other hand, the second group was kept under greenhouse conditions without temperature control, where the temperature fluctuated between 21 and 47 °C, the CO2 levels ranged from 200 to 1300 ppm, and the light intensity varied between 200 and 600 µmol PPFD m−2 s−1. Both groups were maintained in anti-aphid mesh and watered every third day with 500 mL of distilled water for 30 days, and foliar fertilization was applied monthly using 1.5% of a commercial foliar nutrient solution (11-08-06, N-P-K, respectively) (Bayfolan® Forte Liquid, Bayer, Barmen, Germany). For each group, six leaves were measured using a Vernier caliper (with precision of 0.01 mm), and the average value was taken as the leaf size.

2.2. Measurement of Physiological Variables and Data Analysis

Once WP symptoms developed, physiological variables related to chlorophyll, fluorescence, and gas exchange were determined. Ten plants with and without WP symptoms were used. SPAD value obtained was the average of three leaves per plant (three points on each side of chlorotic zone). To determine the normality and homoscedasticity of the data, Shapiro–Wilks and Levene tests were applied, respectively. Then, the obtained data were analyzed using a two-tailed Student’s t-test (p < 0.05) with R software version 4.3.2.

2.3. Chlorophyll and Fluorescence Variables

The SPAD index value was obtained using a SPAD-502 chlorophyll meter (Konica Minolta Sensing, Osaka, Japan). Additionally, to measure the Fv/Fm ratio, as well as the photosynthetic efficiency (PIABS), a multi-function, plant efficiency analyzer (MPEA) chlorophyll fluorometer (FMS 2 Hansatech Instruments Ltd., Norfolk, UK) was employed. The setting conditions were as follows: leaves were adapted for 20 min in darkness, followed by a saturation pulse of 3000 μmol m−2 s−1 (70%) for 1 s. Also, fluorescence variables, such as quantum efficiencies, specific fluxes, phenomenological fluxes, and density of the PSII reaction center were calculated using equations from Strasser et al. [15] (Table 1). Lastly, OJIP curves were constructed using the fluorescence data for both the control plants and those showing WP symptoms.

2.4. Gas Exchange Variables

Net photosynthesis (Pn), stomatal conductance (gs), intercellular CO2 (Ci), and transpiration (E) were measured between 8:00 and 9:00 h using a portable photosynthesis system (LI-COR LI-6400XT Inc., Lincoln, NE, USA). A 6 cm2 chamber was used, and the system was set to a temperature of 25 °C with an airflow rate of 800 μmol s−1, PAR of 800 μmol m−2 s−1, and CO2 flux of 600 μmol s−1.

2.5. Calculation of Water Use and Carboxylation Efficiency

Based on the gas exchange data, Water Use Efficiency (WUE), Intrinsic Water Use Efficiency (IWUE), and Instantaneous Carboxylation Efficiency (ICE) were calculated according to the following formulas:
WUE = Pn/E; IWUE = Pn/gs; ICE = Pn/Ci

3. Results

3.1. Wood Pocket Symptoms and SPAD Index

Persian lime plants, which were not maintained under controlled temperature conditions, exhibited symptoms, such as intense chlorotic mottling and a reduction in leaf size (76.4 ± 15.7 mm × 43.8 ± 9.12 mm) (Figure 1a), consistent with WP symptoms; in contrast, those kept under controlled temperature conditions showed no incidence of WP, and their leaves were larger (111.8 ± 7.2 mm × 58.6 ± 3.2 mm in length) (Figure 1b). Observations under a light stereoscope confirmed the presence of chlorotic areas (Figure 1c), which were absent in healthy leaves (Figure 1d). When correlating the symptoms with chlorophyll content (Figure 1e), it was observed that healthy plants had SPAD index values of 60, while plants showing WP symptoms had SPAD index values close to 30, indicating a statistically significant difference between the two groups (p < 0.05).

3.2. Fluorescence Parameters and OJIP Curves

In healthy plants, the photosynthetic efficiency parameter Fv/Fm remained close to 0.8, while in Persian lime leaves showing WP symptoms, the Fv/Fm value was just above 0.5, indicating a clear level of stress, which results in the loss of energy captured in fluorescence (Figure 2a). On the other hand, the PIABS parameter in healthy plants was close to 8.0, whereas in leaves with WP symptoms, PIABS did not exceed a value of 0.3 (Figure 2b). Both variables showed a significant difference between leaves with and without WP symptoms (p < 0.05). Regarding the OJIP curves, healthy plants exhibited a typical pattern of fluorescence inflection points in each of the phases (O, J, I, P), reaching maximum fluorescence values close to 30,000 relative units (Figure 2c). In contrast, plants showing WP symptoms displayed a completely altered curve; although during the O phase, fluorescence behaved similarly to healthy plants, from the J phase onwards, fluorescence did not increase, resulting in a flattened curve and reaching a maximum fluorescence level of 10,000 relative units, suggesting a significant disruption in the electron transport chain (Figure 2c).
On the other hand, quantum efficiencies in healthy plants reached the following values: TR0/ABS (Fv/Fm) = 0.796, ET0/ABS = 0.495, and ET0/TR0 = 0.621. In contrast, leaves showing WP symptoms exhibited a clear decrease in these efficiencies, with values of TR0/ABS (Fv/Fm) = 0.572, ET0/ABS = 0.157, and ET0/TR0 = 0.250. All three variables were statistically different (p < 0.05). These results indicate that WP directly impacts the plant’s ability to use absorbed energy for electron movement (Table 2).
Regarding specific fluxes, healthy plants showed values of 0.864, 0.839, 0.416, and 0.176 for ABS/RC, TR0/RC, ET0/RC, and DI0/RC, respectively. In contrast, WP causes significant increases (p < 0.05) in the fluxes ABS/RC, TR0/RC, and DI0/RC, reaching values of 3.012, 1.931, and 2.301, respectively, while the ET0/RC flux remained statistically unchanged between healthy plants and those with WP (Table 2).
In terms of phenomenological energy fluxes: trapped energy flux allowing for the reduction of QA per cross-section (TR0/CS0) and electron transport flux per cross-section (ET0/CS0) were higher in healthy control plants than in plants with WP symptoms. However, energy flux per cross-section (ABS/CS0) was higher in plants with WP than in control plants (Table 2).
Lastly, regarding the density of PSII reaction center values before being exposed to actinic light (RC/CS0), healthy plants showed higher values than those showing WP symptoms (p < 0.05). After the exposure to actinic light, the values of density of the PSII reaction center (RC/CSm) increased in both healthy and WP symptomatic plants, but their values were statistically different (p < 0.05), demonstrating that the WP plants show fewer reaction centers in the PSII after the light exposition (Table 2).

3.3. Gas Exchange Parameters and Efficiencies

Physiological variables related to gas exchange show a significant difference (p < 0.05) between healthy plants and those with WP symptoms. First, for net photosynthesis (Pn), healthy plants reached values above 12 µmol CO2 m⁻2 s−1, while plants with WP symptoms barely reached values close to 7 µmol CO2 m⁻2 s−1 (Figure 3a). Additionally, stomatal conductance (gs) in healthy plants remained at 0.1 mol H2O m⁻2 s−1, whereas in plants with WP symptoms, gs reached negative values, indicating that the stomata were tightly closed (Figure 3b). Similarly, transpiration (E) values showed a trend similar to gs, with healthy plants reaching 2.0 mmol H2O m⁻2 s−1, while transpiration values in plants with WP were also negative (Figure 3c). Finally, for intercellular CO2 (Ci), an inverse trend was observed compared to the other variables; healthy plants showed values close to 500 µmol CO2 mol−1, while plants with WP symptoms generated values exceeding 1000 µmol CO2 mol−1 (Figure 3d).
Finally, using the gas exchange variables data, the Water Use Efficiency (WUE) was calculated. In healthy plants, WUE = 5.91 μmol CO2 mmol−1 H2O, whereas in WP plants, the WUE showed negative values of −16.05 μmol CO2 mmol−1 H2O. For the Intrinsic Water Use Efficiency (IWUE), healthy plants displayed high values, exceeding 129 µmol CO2 mol−1 H2O, while plants with WP showed negative values of −150.62 µmol CO2 mol−1 H2O. Lastly, for the Instantaneous Carboxylation Efficiency (ICE), both groups showed positive values but were statistically different (p < 0.05). Healthy plants had an ICE = 0.231 mol m⁻2 s−1, while the WP plants barely reached a value of 0.006 mol m⁻2 s−1. These results are summarized in Table 3.

4. Discussion

The WP physiopathy is a re-emergent condition that has recently captured the attention of Persian lime producers, technicians, and researchers. This physiopathy causes an alteration in the leaves and fruits of Persian lime, leading to decreased quality and economic losses. Despite the importance of WP, very little scientific knowledge is currently available.
In the present study, we show that the symptomatology of WP is associated with a reduction in all physiological parameters evaluated, including a lower SPAD index, higher chlorophyll fluorescence (lower Fv/Fm ratio), and a PIABS lower than that of healthy leaves, indicating damage to PSII, which is one of the first consequences of abiotic stress in plants [16]. In our research, the principal difference between the two groups of plants was the temperature, with symptoms appearing in the group exposed to temperatures over 45 °C. Previously, the exposure of citrus plants to high temperatures has been associated with PSII damage, loss of redox balance, significant foliar damage [17], and the degradation of citric acid [18]. We observed that, in leaves with WP symptoms, the transpiration (E) and the stomatal conductance (gs) were negative, indicating closure of stomata, impacting in the WUE and IWUE, contrary to what is observed in high-temperature-tolerant citrus genotypes, where the stomatal fluxes are increased due to a lower accumulation of abscisic acid (ABA) [19]. In this study, a reduction in photosynthesis (Pn) was clearly observed in leaves with WP symptoms, which is consistent with previously published studies of citrus genotypes exposed to high temperatures [17,20,21].
One of the principal effects of high-temperature stress in plants is the increase in Reactive Oxygen Species (ROS) in cells, which leads to the activation of photorespiration and is associated with increased oxidative stress, degradation of chloroplasts and photosynthetic pigments, and reduced efficiency of energy use by PSII [16]. The photorespiration process is a mechanism by which the plant uses the Rubisco enzyme to metabolize oxygen instead of carbon dioxide, recycling the toxic 2-phosphoglycolate (2PG) into 3-phosphoglycerate (3PG) and reintroducing it into the Calvin–Benson cycle [22]. However, the photorespiration process is highly complex and wasteful due to its use of ATP [23]. As a result, photorespiration inhibits carboxylation, leads to the accumulation of intercellular CO2, and reduces photosynthesis in plants [22], which can be triggered by environmental factors, such as high temperatures in citrus [24], a result observed in our study.
Recently, it has been demonstrated that inhibition of photorespiration through metabolic engineering [25] and the use of alternative electron acceptors [26] confers tolerance to high-temperature stress. Thus, we hypothesize that exposure of Persian lime to high temperatures could generate an exacerbated photorespiration process, which causes a reduction in stomatal fluxes and impacts plant homeostasis, leading to the appearance of WP symptoms. This hypothesis is consistent with observations of recent warming of the regions where Persian lime is currently cultivated [27].
WP physiopathy has been associated exclusively with Persian lime, a triploid citrus with genetic heritage from all four true citrus species [28], making the physiology of this species highly complex. For example, Persian lime is highly tolerant to Huanglongbing disease due to its ability to maintain stomatal fluxes during infection [29]. Additionally, water availability during specific phenological stages is crucial for yield and fruit quality, as it is linked to stomatal conductance [8]. Similarly, other triploid citrus genotypes have been reported to tolerate chilling, associated with the maintenance of photosynthesis and stomatal conductance [30]. Higher stomatal fluxes are markers of abiotic stress alleviation in triploid citrus; however, in WP-affected Persian lime plants, the stomata remain closed. The closure of stomata could be orchestrated by the accumulation of oxidative markers, such as H2O2 and MDA, leading to the overproduction of ABA [30,31,32,33]. In this context, Trueba et al. [34] reported that severe drought causes important physiological responses, such as stomatal closure, which in turn affects transpiration and photosynthesis; the magnitude of the reaction depends on water-deficit stress. Likewise, in Tahiti acid lime, irrigation and nutrition management allowed for improving the functional and physiological response of trees with phisiopathy, associating such a condition with the increase in stomatal conductance and photosynthesis, which, in turn, increased the average yield and renewed the trees foliage [35].
In plants, increased ABA levels are inversely proportional to chlorophyll levels [36,37] and inhibit the transcription of chloroplast genes [38]. Thus, our results suggest that elevated ABA levels in Persian lime leaves, potentially triggered by high-temperature stress, cause stomatal closure and degradation of photosynthetic pigments, leading to the characteristic WP symptomatology. However, this hypothesis must be corroborated experimentally.

5. Conclusions

In this study, we present the first physiological characterization of WP physiopathy in Persian lime that shows that symptomatic leaves exhibit alterations in all physiological parameters evaluated, including fluorescence and gas exchange variables. Based on the physiological data, we propose two hypotheses: (1) high-temperature stress in Persian lime leaves exacerbates the photorespiration process due to stomatal closure, and (2) that stomatal closure is associated with an accumulation of ABA levels in response to high-temperature stress. However, further studies are needed to evaluate these hypotheses and design effective control strategies.

Author Contributions

Conceptualization, F.R.F.-d.l.R., R.S.-M., H.E.-M. and J.M.S.; methodology, F.R.F.-d.l.R., G.F.-O. and C.M.-H.; formal analysis, F.R.F.-d.l.R., R.S.-M. and H.E.-M.; investigation, F.R.F.-d.l.R., R.S.-M., H.E.-M. and J.M.S.; data curation, F.R.F.-d.l.R. and G.F.-O.; writing—original draft preparation, F.R.F.-d.l.R. and R.S.-M.; writing—review and editing, F.R.F.-d.l.R., R.S.-M., C.M.-H. and H.E.-M.; visualization, F.R.F.-d.l.R.; supervision, J.M.S.; funding acquisition, H.E.-M. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the Consejo de Productores y Exportadores de Limón Persa, A.C. (COPELP), through the project: “Mapeo regional y caracterización genética de la fisiopatía denominada Wood Pocket en limón Persa de la región citrícola de Martínez de la Torre, Ver. y Huimanguillo, Tab.” (No. 13415836195).

Data Availability Statement

The original contributions presented in this study are included in the article. Further inquiries can be directed to the corresponding authors.

Acknowledgments

The authors would like to thank the Eduardo Castillo-Castro and Nelly González-Oviedo, for the technical support in the physiological analysis.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. Comparison between healthy (b,d) and WP symptomatic leaves (a,c). SPAD index difference between healthy and WP symptomatic leaves (e). Whiskers indicate ± SD. Asterisks represent statistical difference according to Student’s t-Test (p < 0.05).
Figure 1. Comparison between healthy (b,d) and WP symptomatic leaves (a,c). SPAD index difference between healthy and WP symptomatic leaves (e). Whiskers indicate ± SD. Asterisks represent statistical difference according to Student’s t-Test (p < 0.05).
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Figure 2. Fluorescence parameters evaluated in healthy and WP symptomatic leaves of Persian lime: (a) Fv/Fm ratio, (b) PIABS, and (c) OJIP curves. Whiskers represent ± SD. Asterisks represent statistical difference according to Student’s t-Test (p < 0.05).
Figure 2. Fluorescence parameters evaluated in healthy and WP symptomatic leaves of Persian lime: (a) Fv/Fm ratio, (b) PIABS, and (c) OJIP curves. Whiskers represent ± SD. Asterisks represent statistical difference according to Student’s t-Test (p < 0.05).
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Figure 3. Gas exchange parameters measured in Persian lime leaves with and without WP symptoms: (a) photosynthesis net (Pn), (b) stomatal conductance (gs), (c) transpiration (E), and (d) intercellular carbon (Ci). Whiskers indicate ± SD. Asterisks represent statistical difference according to Student’s t-Test (p < 0.05).
Figure 3. Gas exchange parameters measured in Persian lime leaves with and without WP symptoms: (a) photosynthesis net (Pn), (b) stomatal conductance (gs), (c) transpiration (E), and (d) intercellular carbon (Ci). Whiskers indicate ± SD. Asterisks represent statistical difference according to Student’s t-Test (p < 0.05).
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Table 1. Summary of technical fluorescence parameters obtained from the fast Chl fluorescence transient according to Strasser et al. [15]. ABS: absorption energy flux, CS: excited cross-section of leaf sample, ET: flux of electrons from QA into the electron transport chain, RC: reaction center of PSII, TR: excitation energy flux trapped by a RC and utilized for the reduction of QA to QA−.
Table 1. Summary of technical fluorescence parameters obtained from the fast Chl fluorescence transient according to Strasser et al. [15]. ABS: absorption energy flux, CS: excited cross-section of leaf sample, ET: flux of electrons from QA into the electron transport chain, RC: reaction center of PSII, TR: excitation energy flux trapped by a RC and utilized for the reduction of QA to QA−.
Technical Fluorescence Parameters
Quantum efficiencies
TR0/ABS=Fv/FmMaximum quantum yield for primary photochemistry.
ET0/ABS=Fv/Fm × (1 − Vj)Probability that an absorbed photon will move an electron into electron transport chain further than QA−.
ET0/TR0=(1 − Vj)Efficiency by which a trapped exciton, having triggered the reduction of QA to QA−, can move an electron further than QA− into the electron transport chain.
Specific fluxes
ABS/RC=M0 × (1/Vj) × [1/(TR0/ABS)]Chlorophyll antenna’s absorption flux per RC.
TR0/RC=M0 × (1/Vj)Trapped energy flux that leads to QA reduction per RC.
ET0/RC=M0 × (1/Vj) × (ET0/TR0)Electron transport flux, further than QA, per RC.
Phenomenological fluxes
ABS/CS0=ABS/CSChl =Chl/CS0 or ABS/CS0 = F0 or ABS/CSm = Fm Absorbed energy flux per excited cross-section of leaf sample at F0.
TR0/CS0=ABS/CS0 × TR0/ABSTrapped energy flux that leads to QA reduction per excited cross-section of leaf sample.
ET0/CS0=ABS/CS0 × (TR0/ABS) × (ET0/TR0)Electron transport flux (further than QA) per excited cross-section of leaf sample.
Density of PSII reaction center
RC/CS0=(ET0/TR0) × (Vj/M0) × (ABS/CS0)Reaction center per cross-section at F0.
RC/CSm=(ET0/TR0) × (Vj/M0) × (ABS/CSm)Reaction center per cross-section at Fm.
Table 2. Measured fluorescence variables for leaves of Persian lime plants, with and without WP symptoms. Data are means ± SD (n = 10). Different letters mean statistical differences according to Student’s t-test (p < 0.05).
Table 2. Measured fluorescence variables for leaves of Persian lime plants, with and without WP symptoms. Data are means ± SD (n = 10). Different letters mean statistical differences according to Student’s t-test (p < 0.05).
ControlWood Pocket
Quantum efficiencies
TR0/ABS 0.796 ± 0.012 a0.572 ± 0.134 b
ET0/ABS0.495 ± 0.069 a0.157 ± 0.105 b
ET0/TR00.621 ± 0.084 a0.250 ± 0.147 b
Specific fluxes
ABS/RC0.864 ± 0.058 a3.012 ± 0.771 b
TR0/RC0.839 ± 0.374 a1.931 ± 0.350 b
ET0/RC0.416 ± 0.074 a0.443 ± 0.220 a
DI0/RC0.176 ± 0.021 a2.301 ± 1.810 b
Phenomenological fluxes
ABS/CS05808.83 ± 301.95 a6683.25 ± 1068.03 b
TR0/CS04620.65 ± 183.27 a4399.00 ± 1628.54 a
ET0/CS02878.56 ± 419.70 a1325.13 ± 755.22 b
Density of PSII reaction center
RC/CS04724.44 ± 301.67 a1137.99 ± 898.57 b
RC/CSm25,536.79 ± 3985.89 a3928.65 ± 1820.40 b
Table 3. Efficiency calculations based on gas exchange parameters of Persian lime plants with and without Wood Pocket symptoms. Data are means + standard deviation (n = 5). Different letters mean statistical differences according to Student’s t-test (p < 0.05).
Table 3. Efficiency calculations based on gas exchange parameters of Persian lime plants with and without Wood Pocket symptoms. Data are means + standard deviation (n = 5). Different letters mean statistical differences according to Student’s t-test (p < 0.05).
TreatmentWater Use Efficiency
(μmol CO2 mmol−1 H2O)		Intrinsic Water Use Efficiency
(µmol CO2 mol−1 H2O)		Instantaneous Carboxylation Efficiency
(mol m−2 s−1)
Healthy5.91 ± 1.17 a129.08 ± 8.59 a0.0231 ± 0.003 a
Wood Pocket−16.05 ± 5.87 b−150.62 ± 21.42 b0.006 ± 0.001 b
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Flores-de la Rosa, F.R.; Fuentes-Ortíz, G.; Santillán-Mendoza, R.; Matilde-Hernández, C.; Estrella-Maldonado, H.; Santamaría, J.M. Understanding the Wood Pocket Physiopathy in Persian Lime Through Its Physiological Characterization. Agronomy 2025, 15, 762. https://doi.org/10.3390/agronomy15040762

AMA Style

Flores-de la Rosa FR, Fuentes-Ortíz G, Santillán-Mendoza R, Matilde-Hernández C, Estrella-Maldonado H, Santamaría JM. Understanding the Wood Pocket Physiopathy in Persian Lime Through Its Physiological Characterization. Agronomy. 2025; 15(4):762. https://doi.org/10.3390/agronomy15040762

Chicago/Turabian Style

Flores-de la Rosa, Felipe Roberto, Gabriela Fuentes-Ortíz, Ricardo Santillán-Mendoza, Cristian Matilde-Hernández, Humberto Estrella-Maldonado, and Jorge M. Santamaría. 2025. "Understanding the Wood Pocket Physiopathy in Persian Lime Through Its Physiological Characterization" Agronomy 15, no. 4: 762. https://doi.org/10.3390/agronomy15040762

APA Style

Flores-de la Rosa, F. R., Fuentes-Ortíz, G., Santillán-Mendoza, R., Matilde-Hernández, C., Estrella-Maldonado, H., & Santamaría, J. M. (2025). Understanding the Wood Pocket Physiopathy in Persian Lime Through Its Physiological Characterization. Agronomy, 15(4), 762. https://doi.org/10.3390/agronomy15040762

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