Temperature-Driven Variability in Melon Root System Architecture

Temperature-Driven Variability in Melon Root System Architecture | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Article Temperature-Driven Variability in Melon Root System Architecture Amnon Cochavi, Elad Oren, Fabian Baumkoler, Evyatar Asaf, Evgeny Smirnov, and 2 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8126587/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 13 You are reading this latest preprint version Abstract Shifting the sowing date of summer crops such as melon ( Cucumis melo L.) to early spring presents both agronomic opportunities and challenges. Under suboptimal temperatures, slowed physiological development allows weeds to gain a competitive advantage. Enhancing early seedling vigor under cool conditions is therefore essential for successful crop establishment and effective weed suppression. In this study, ten melon accessions representing a broader collection (~ 180) were evaluated for early vigor and performance under suboptimal temperatures. Pulling resistance, root system scanning, and physiological measurements were used to identify key traits associated with early root establishment. Accessions were compared under optimal (25°C) and suboptimal (16°C) temperature regimes. Pulling strength showed a strong correlation with root biomass and volume across accessions. The AY accession exhibited clear tolerance to low temperatures, maintaining high physiological activity, whereas PI414723 displayed reduced carbon assimilation due to diffusional limitations. These findings demonstrate that tolerance to suboptimal temperatures exists even within warm-climate crops such as melon and highlight the potential of AY as a valuable genetic resource for developing early-season, stress-tolerant melon varieties. Biological sciences/Ecology Earth and environmental sciences/Ecology Biological sciences/Physiology Biological sciences/Plant sciences diffusional limitation pulling strength root traits suboptimal temperatures Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Introduction In response to climate change, many melon ( Cucumis melo L.) growing regions have shifted sowing from the traditional summer season to late winter or early spring to reduce the adverse effects of elevated temperatures and the associated increase in evapotranspiration and pest pressure. 1,2 . However, early-season planting under suboptimal temperatures slows crop development, and therefore promotes weed dominance, and complicates mechanical weeding 3 . These challenges highlight the need to identify melon accessions with rapid early-stage growth and adaptation to suboptimal temperatures Melon is one of the most popular fruits globally ( http://faostat3.fao.org/ ). Its origin traces back to warm regions in Central and Southern Africa, with related species found in other warm areas, including the Indian peninsula, Asia, and Australia 4,5 . Consequently, melon domestication occurred independently in multiple regions 5 , leading to a diverse range of plant traits 6,7 . This variety provides a valuable resource for improving melon adaptation to different agro-technical and climatic growth conditions. Weed control is a major challenge in open-field melon cultivation 8 . Melon is a weak competitor against weeds and experiences significant yield losses under high weed infestation levels 9 . Purple nutsedge (Cyperus rotundus), Puncturevine ( Tribulus terrestris L.), Johnsongrass ( Sorghum halepense ), and pigweeds ( Amaranthus spp.), are just some of the summer weeds that negatively affect the growth and yield of melon. The melon’s horizontal growth habit, characterized by long, trailing vines, often fails to achieve full canopy closure through the growing season, leaving the soil exposed to light. This exposure promotes multiple weed emergence flushes and reduces melon’s ability to suppress weed growth 9 . Furthermore, as with many minor crops, registered herbicide options for melon are limited, providing only partial weed control. Thus, mechanical weeding remains the most effective non-chemical mean for weed removal in melon fields 8,10 . For mechanical weeding to be effective and safe for the crop, melon plants must have stronger resistant to pulling by the mechanical device than competing weeds. Crop plants with early vigor have a competitive advantage against weeds 11 . Early and rapid canopy establishment suppresses weed growth by limiting their access to light 12 . In addition to the above-ground suppression, a well-developed root system can hinder weed emergence and establishment 13 . Strong root development also enhances plant resilience to environmental fluctuations, resource limitations, and various biotic and abiotic stresses 14–17 . Most early vigor studies determining differences among various varieties and/or accessions focused on wheat ( Triticum sp.) and other cereal crops (Aharon et al., 2021 ), while research on dicot crops remains limited 18 . To our knowledge, no previous study has examined the diversity of melon accessions for their early vigor under varying temperatures. Early vigor is strongly affected by environmental factors, particularly temperature, which drives genotype-by-environment (G × E) interactions. Low and suboptimal temperatures are well-documented to inhibit physiological, enzymatic, and molecular processes in plants 19–21 . Thus, identifying melon accessions with consistent root development under moderate and suboptimal temperatures has multiple advantages. It can enhance early vigor and weed competitiveness across varying climates while also ensuring safe and effective mechanical weed control. In turn, this approach could contribute to more sustainable weed management strategies in melon cultivation. Root system phenotyping is challenging due to its belowground development and the destructive nature of most measurement methods 22,23 . Therefore, field-based root assessments and screening breeding populations for root traits require intensive labor. In this context, root pulling using a digital force gauge provides a simple and effective first line method for assessing root system pulling strength under both field and laboratory conditions 10 . Given the link between pulling strength and root system structure, this method serves as a useful proxy for root system dynamics under varying conditions 24–26 . The force required to uproot a plant is strongly correlated with key root traits such as length, diameter, biomass and other parameters 27–29 . However, this method should be used cautiously to minimize biases introduced by soil properties (e.g., texture, soil water content) and pulling direction 29 . Suboptimal temperatures significantly inhibit plant physiological activity. Previous studies have identified the minimum, optimum, and maximum temperatures for melon growth as 10°C, 34°C, and 45°C, respectively 30 . These conditions reduce chlorophyll production, decrease membrane fluidity, and restrict nutrient translocation, ultimately limiting plant function 20,21,31 . Warm-region crops, such as melons, are sensitive to moderate temperature reductions and brief frost events 32 . During abiotic stress, such as unfavorable temperatures, the translocation of carbohydrates and minerals from roots to shoots can 33–36 . Therefore, physiological measurements can provide insights into differential responses among accessions, aiding in the identification of those with adaptive traits. The combined impact of suboptimal temperatures and weed competition can drastically reduce crop productivity 37 . While weed competitiveness remains high across temperature ranges, crop development, especially in warm-region species like melon, is significantly impaired under suboptimal temperatures. At early growth stages, the rapid expansion of weeds alters the red:far-red (R:Fr) light ratio, suppressing both above- and below-ground development of the crop. This effect can persist long after weeds are removed, leading to lasting yield reductions 38 . Consequently, effective and timely weed control under suboptimal temperatures is critical to preventing long-term damage and ensuring stable crop yields. Current knowledge of melon root system diversity and function remains limited, and the relationship between root system architecture and mechanical pulling strength is still poorly understood. Only a few studies have examined variation among melon accessions in these traits. To develop sustainable strategies for mechanical weed management during early melon growth, a deeper understanding of root system characteristics and their mechanical properties is therefore essential. This study aims to screen and characterize a diverse melon population for early development, with a particular focus on performance under suboptimal temperature conditions. By integrating the pulling method, root system screening, and leaf-level gas exchange and chlorophyll fluorescence measurements, the study seeks to identify melon accessions combining tolerance to suboptimal temperatures with strong and resilient root systems. Materials and methods Plant material and growing conditions From the Newe Ya'ar collection of 177 melon ( Cucumis melo L.) accessions (Gur et al., 2017 ), 10 accessions representing the main variety types were selected (Table 1 ). Plants were grown in 2-liter pots (Pot dimensions: top diameter = 16 cm, base diameter = 14 cm, height = 14 cm; volume ≈ 2.46 l) filled with local soil (chromic Haploxerert, fine-clayey, montmorillonitic, thermic) comprising 55% clay, 25% silt, and 20% sand. Soil porosity was found to be ~ 60%, the bulk density of the soil was 1.02 g cm − 3 , where field capacity and wilting point is 37% and 22% respectively (see appendix 1). The final mixture contained 2% organic matter and a pH of 7.2 (see appendix 1). Each accession was grown in five biological replicates. Table 1 The ten representative melon accessions used in this work from the Newe Ya’ar collection. The accessions are not subject to patent protection. Line Name Variety Name Sub-Species varietyType Marketing Type Origin (Company) AY Ananas Yoqne'am melo Reticulatus Ananas Israel (H’azera) DUL Dulce melo Reticulatus American cantaloupe USA (Bonanza) NA Noy Amid melo Inodorus Yellow Canary Israel (Newe Ya’ar) NY Noy Yizre'el melo Cantalupensis Ha’Ogen Israel (Newe Ya’ar) PI414723 PI414723 agrestis Momordica N.D India (Pitrat M. - INRA) PSR Piel De Sapo Redon melo Inodorus Piel De Sapo Spain (DEFO) QME Qishu Meshulash C. callosus N.D N.D Israel (Wild accession) SAS Sakata Sweet (17–725) agrestis Makuwa N.D Japan (Skata) TAD TAM DEW melo Inodorus Honey Dew USA (Texas A&M) VEP Vedrantais melo Cantalupensis Charentais France (Pitrat M. - INRA) N.D. - Not Defined Table 1 . Linear correlation coefficients between days after planting and plant anchoring strength under two temperature regimes. 25 16 Accession Slope Intercept r 2 Slope Intercept r 2 AY 0.5 2.82- 0.96 0.53 4.4- 0.95 NY 0.72 7.70- 0.96 0.71 15.66- 0.99 Dulce 0.68 6.72- 0.86 0.44 8.4- 0.92 PI414723 0.79 11.08- 1.00 0.12 1.12- 0.98 Plants pulling In the first experiment, the 10 accessions from the Newe Ya'ar collection were grown under greenhouse conditions during the winter of 2022 (planting date - January 3rd ) under natural light conditions. Temperature was set to a minimum of 18ºC (heating activated) and a maximum of 30 ºC (fan activated) with an average temperature of ~ 20 ºC. Plants were pulled at three phenological stages: two to three true leaves, three to four true leaves, and four to five true leaves. Each measurement point (stage by accession) had five replicates. Pulling strength was measured using a digital force gauge (AXIS, FB500N, range 0.1–50 N). The plant shoots were attached to the device with a plastic clamp and pulled horizontally, recording the maximal pulling strength for each plant. To ensure uniform soil properties, all plants were irrigated to field capacity on the morning of the pulling day, and pulled after two hours when water level is still under field capacity (figure S1 ). Soil water content was measured using a true time-domain reflectometer (TDR; Acclima Inc.) following a two-point calibration (0 and 100%) conducted weekly. After the third pulling event, the soil was washed gently with water, and the root system was scanned and analyzed using WinRhizo (Regent Instruments Inc., Ottawa, ON, Canada). Roots were scan in high resulotion of 600 DPI, to ensure the detection of small diameter roots. Roots and the shoots were then oven-dried for 48 hours at 72ºC before weighing. Then, the specific root length (SRL, g cm − 1 ) and specific root surface area (SRA, g cm − 2 ) were calculated from the WinRhizo scanning and dry weight measurement. Four accessions different in their pulling strengths were selected to represent the variety in the collection. These accessions were grown in controlled environment rooms under two temperature regimes: low (16 ºC) and ambient (25 ºC). Plants were grown under artificial photosynthetic active radiation (PAR, 400–700 nm) lights (~ 1000 µmol m − 2 s − 1 , 12/12 hours day/night period), simulating early spring daylight conditions. Pulling was performed at three developmental stages, as outlined in the first experiment, with five replicates for each accession at each stage, totaling 120 plants. Roots and shoots were then dried and weighed as described above. From the four selected accessions, two with distinct temperature responses were chosen for further study. These accessions were grown under identical controlled environmental conditions as described previously. At the four to five true leaf stage, physiological measurements, including CO₂ and H₂O fluxes, along with chlorophyll fluorescence, were conducted using the LI-COR 6800F system (Licor, Lincoln, NE, US). Pulling strength measurement, root scanning, and below and above biomass evaluation were conducted as described above, with five replicates per treatment. Gas exchange and fluorescence parameters In the second controlled environment experiment, gas exchange and fluorescence were measured using the Li-6800F system. Measurement conditions were set to 415 ppm CO₂, 40% relative humidity, a light intensity of 1000 µmol m⁻² s⁻¹, and a temperature corresponding to the ambient room temperature. The youngest fully matured leaf was measured in all plants. Photosynthetic efficiency was assessed using the Pulse Amplitude Modulation (PAM) method, which was conducted simultaneously with flux measurements. To evaluate dark-adapted PAM parameters, measurements were taken in the early morning before the lights were turned on. The functionality of PSⅡ ( \(\:\frac{{F}_{v}}{{F}_{m}}\) ; Sipka et al., 2021 ) was evaluated using the following equation: \(\:\frac{{F}_{v}}{{F}_{m}}=\frac{{F}_{m}-{F}_{0}}{{F}_{m}}\) [1] Where \(\:{F}_{m}\) and \(\:{F}_{0}\) are the maximal and minimal fluorescence measured under dark conditions. The light acclimated quantum yield of PSⅡ (Y(Ⅱ)) was calculated as: \(\:Y\left(Ⅱ\right)=\frac{{F}_{m}^{{\prime\:}}-\:{F}_{0}^{{\prime\:}}}{{F}_{m}^{{\prime\:}}}\) [2] Where \(\:{F}_{m}^{{\prime\:}}\) and \(\:{F}_{0}^{{\prime\:}}\) are the maximal and minimal fluorescence measured under light conditions. Statistical analysis Data normality was assessed using the Shapiro-Wilk test, and homogeneity of variance was tested using Levene's test. Differences between accessions were analyzed using one-way analysis of variance (ANOVA), followed by Tukey-HSD with Bonferroni correction for multiple comparisons (α < 0.0.05). Results Variation in pulling strength indicates differences in root morphology In the initial screening of 10 representative melon accessions, we assessed early vigor and variation in the plant pulling force. No significant differences in pulling strength were observed among accessions at the first two phenological stages (two to four true leaves). However, at the third stage (four to five true leaves), distinct responses emerged (Figure S2). The AY and TAD accessions exhibited the highest anchoring strength, 11.52 and 9.92 N, respectively. In contrast, both the wild (QME) and commercial accessions (DUL) had the lowest values, 1.72 and 1.8 N, respectively (Fig. 1 ). No significant effect of accession, subspecies, or variety was found on pulling strength. In addition to the plant pulling strength, we evaluated the root traits of each melon accession. A strong correlation was observed between root weight and volume in all plant accessions (Fig. 2 a). For example, root volumes of AY (high pulling strength) and QME (low pulling strength) were 4.25 and 2.38 cm 3 , respectively. However, all 10 accessions maintained a similar root-to-shoot ratio (Fig. 2 b). Correlation analysis of all the measured components (root traits, weight, and their derivatives) with the pulling strength revealed that root weight (R 2 = 0.77) and root volume (R 2 = 0.83) were the strongest predictors of pulling strength, while specific root volume showed a weak correlation (R 2 = 0.21; Fig. 3 ). Moreover, all accessions exhibited a consistent relationship between pulling strength, root volume, and weight under these conditions (Fig. 4 A-B). T emperature-driven variation in pulling strength and biomass across accessions In order to confirm the results of the pulling experiment, four accessions with different pulling strengths: the highest (AY), the lowest (Dulce), and two lines with moderate pulling strength (PI414723, and NY). The accessions were grown under two temperature regimes 16º and 25º. The pulling and weight results revealed significant variation in accession responses to temperature (Figs. 5 – 6 , Tables 2 –3). Table 2 Linear correlation coefficients between days after planting and plant shoot biomass under two temperature regimes. 25 16 Accession Slope Intercept r 2 Slope Intercept r 2 AY 0.09 -0.80 0.97 0.1 1.31- 0.98 NY 0.13 1.81- 0.99 0.14 3.36- 0.87 Dulce 0.23 4.17- 0.90 0.13 3.26- 0.83 PI414723 0.24 4.59- 0.92 0.0011 -0.24 0.90 The AY accession demonstrated the highest tolerance to suboptimal temperatures, reaching the two- to three true leaves stage 10 days earlier than the other accessions (Fig. 5 A, 6 A). Biomass accumulation rates remained stable at both 16°C and 25°C, with 0.1 and 0.09 g day − 1 . Although NY emerged later than AY, it also exhibited high stability under both temperature conditions. Pulling strength rates were 0.72 N day − 1 (16°C) and 0.71 N day − 1 (25°C), while biomass accumulation rates were 0.13 g day − 1 (16°C) and 0.14 g day − 1 (25°C), even surpassing AY development rates. In contrast, Dulce and PI414723 accessions were highly susceptible to suboptimal temperatures. While under 25ºC, both accessions demonstrated the highest biomass accumulation rates (0.23 g day − 1 for Dulce, 0.24 g day − 1 for PI414723). However, under suboptimal temperatures, their performances declined significantly, with Dulce’s biomass accumulation reduced by ~ 40%, while PI414723’s growth was nearly eliminated (~ 100% reduction). The root system response is related to the physiological response To further investigate temperature response, we selected two accessions with different tolerances: stable (AY) and susceptible to suboptimal temperatures (PI414723). The AY accession was grown with five replicates per temperature, whereas PI414723 was grown with four replicates due to seedling collapse. Across both accessions, all growth parameters declined under suboptimal temperatures (Fig. 7 ). The difference in pulling strength between the tolerant and susceptible accessions was minor at 25ºC (7.6 N for AY and 11.2 N for PI414723) but became pronounced at 16ºC (6.1 N for AY and 2.8 N for PI414723; Fig. 7 a). Root biomass did not significantly differ between the accessions across temperatures (Fig. 7 b). However, at 25ºC, AY exhibited significantly greater root length (1210 cm) and root volume (5.62 cm³) compared to PI414723, which had lower values at both 25ºC (267 cm and 0.62 cm³) and 16ºC (214 cm and 0.59 cm³). Physiological measurements further supported these findings (Fig. 8 ). Carbon assimilation rates declined in both accessions under suboptimal temperatures (under 16ºC). However, while AY showed a 40% reduction, PI414723 experienced a 70% reduction (Fig. 8 a). Stomatal conductance remained constant in AY, whereas PI414723 showed a 90% reduction at 16 ºC (Fig. 8 b). Lastly, PSⅡ functionality ( \(\:\frac{{F}_{v}}{{F}_{m}}\) ) decreased in both accessions, 25% in AY and 50% in PI414723 (Fig. 8 c). The PSⅡ quantum yield (Y(Ⅱ)) was reduced by 50% in AY and 65% in PI414723 under suboptimal conditions (Fig. 8 d). Discussion The historical journey of melon ( Cucumis melo ) from its origins in the arid regions of Africa, through the Indian subcontinent, to East Asia and eventually the Western world, has greatly enhanced its internal diversity, ranging from physiological adaptations to the wide variability in fruit traits 5,7,40 . Both natural selection and human-mediated cultivation across diverse latitudes have contributed to this variability, increasing the likelihood of identifying accessions within the species that possess tolerance to both heat and cold stress 41 . The results of the present study highlight that even in a warm-season crop such as melon, accessions exhibiting tolerance to suboptimal temperatures can be found, likely as a result of indirect selection. The AY accession, for instance, was originally cultivated on specially prepared fallow fields under dryland conditions without irrigation, sown in early spring, and relied entirely on soil moisture conserved from winter precipitation 42 . This early sowing during late winter or early spring suggests that this accession possesses physiological mechanisms that enable tolerance to such unfavorable environmental conditions. Moreover, its ability to rely on residual soil moisture from winter precipitation implies the presence of a strong and efficient root system, capable of sustaining growth under limited water availability. Stomata closure is one of the first line process in response to temperature decrease 21,43 . This in turn, led to a reduction on the photosynthetic activity and decrease in the carbon assimilation rate 31,44 . Under optimal conditions, both accessions exhibited similar physiological behavior. However, under suboptimal conditions, their responses were distinct. While AY stomata remain open under suboptimal temperatures, the PI414723 stomatal conductance decrease sharply under same conditions. This stomatal closure induced additional feedback inhibition of photosynthesis and therefore on the carbon assimilation rate 45 . In addition, Zait et al., ( 2024 ) demonstrated that fluorescence parameters exhibit a gradual response to stress, with \(\:\frac{{F}_{v}}{{F}_{m}}\) reduction serving as a late-stage indicator, occurring at higher stress levels. At this stage, the decrease in the xanthophyll pool may mask other parameters, such as Y(II). This can explain the similar Y(II) values observed in both accessions under suboptimal temperatures, although PI414723 was under greater stress under these conditions. Several pathways can lead to stomata closure under suboptimal conditions as abscisic acid (ABA) and calcium movement in the plant 31,43 . In the current work, the AY stomata seems to be inert to the temperature reduction. This accession, cultivated for more than 100 years, challenge our understanding if it was inbreed as suboptimal temperatures accession for early sowing, or it was selected for this trait over the last decades 7,42 . The limited response to ABA can also explain the early emerge of the AY seedlings under sub optimal temperatures. Tolerance to suboptimal temperatures can be expressed in both root and shoot tissues, with physiological and molecular adjustments in one organ potentially influencing the other through systemic signaling and resource allocation 47,48 . Such cross-organ interactions may involve hormonal and molecular regulation, metabolic adjustments, and shifts in source–sink dynamics that collectively contribute to whole-plant acclimation under suboptimal temperature conditions 49,50 . The current study demonstrated a direct link between aboveground physiological responses and belowground development under suboptimal temperature. In the tolerant AY accession, both root and shoot growth rates remained stable across low and ambient temperature conditions. In contrast, the susceptible PI414723 accession exhibited a concurrent reduction in root and shoot development, consistent with a marked decline in key physiological parameters. Early crop vigor is a critical trait for early-season sowing, as it enhances weed suppression and promotes successful crop establishment 11 . Furthermore, genotype × environment (G × E) stability is essential to ensure consistent crop performance under the variable environmental conditions that occur throughout the growing season 32,34 . The importance of maintaining such stability is becoming increasingly evident in the context of climate change, where crops must perform reliably under fluctuating and often stressful conditions. This challenge is further intensified by the global shift toward reducing chemical inputs in agricultural systems, which demands varieties with inherently robust and adaptive traits 51 . Although the AY accession did not exhibit the fastest development under optimal conditions, it maintained a stable developmental rate across both temperature regimes. This stability, combined with its 10-day developmental advantage under suboptimal temperatures, likely contributes to its suitability as an early-season cultivar with both agronomic and economic value. Early emergence enhances crop competitiveness and management efficiency, while also providing farmers with clear economic benefits through improved resource utilization and earlier market access. Precise and highly accurate root system phenotyping methods are often complex, costly, and require specialized equipment, which limits their accessibility and scalability for large surveys. For instance, several studies have employed X-ray computed tomography (CT; Lobet & Draye, 2013 ; Rellan- Alvarez et al., 2015 ), or custom-designed growth boxes 54 to investigate belowground dynamics. While these approaches provide valuable insights into root architecture and function, their major limitation lies in their unsuitability for large-scale screening, which is essential for evaluating extensive plant populations such as those examined in the present study. Despite its simplicity, the pulling method employed in this study showed a strong and consistent correlation with key root traits, particularly root volume and biomass, across all melon accessions and temperature regimes. Similar relationships between pulling strength and root system characteristics, such as total root length and volume, have also been reported in previous studies 28,29 . Therefore, this method represents a practical and effective tool for large-scale, first-step screening of plant populations or for field validation of more advanced and resource-demanding root phenotyping approaches. It is important to recognize that the relevance of the measured root traits extends beyond mechanical weed management, as they also play a central role in nutrient and water uptake 15,55 . Consequently, these traits can strongly influence plant responses to nutrient limitation and water scarcity. Future studies on root system behavior could benefit from integrating this simple pulling method with more precise phenotyping approaches, thereby enabling a more comprehensive understanding of the functional and structural characteristics of root systems under varying environmental conditions. In the present study, we successfully characterized root system responses in a large melon population using the pulling method, which was validated through root system scanning. The measurement device, a simple force gauge, is affordable, widely accessible, and easy to operate without the need for specialized training. Moreover, the method is time-efficient and provides a reliable first-line assessment of key root traits. Applying this approach as an initial screening tool prior to more labor- and time-intensive analyses could substantially improve the efficiency of studies on root dynamics and physiology. Future research on these traits is supported by the availability of fully de novo assembled and annotated genomes for the melon accessions evaluated in this study, including AY, along with readily available segregating populations 7 . These genomic resources provide a strong foundation for investigating the genetic basis of root pulling resistance, early seedling vigor, and temperature tolerance. Integrating these physiological and genomic insights could ultimately enable the development of melon varieties or specialized rootstocks optimized for sustainable, non-chemical weed management. Such innovations would enhance melon adaptability to both current and future environmental conditions, contributing to more resilient and resource-efficient agricultural systems. Conclusions The current study demonstrated that the considerable genetic diversity within this warm-climate crop results in a wide range of responses to temperature. The limited negative response to suboptimal temperatures observed in the tolerant AY accession led to earlier emergence and higher physiological activity compared to other melon accessions. This likely explains its longstanding use as an early-season cultivar. The combination of rapid emergence and vigorous root system development enables its growth without supplemental irrigation, making it well-suited for conditions that favor early-season weed competition and mechanical weeding. Although the fruits of this accession are not compatible with current market demands, mainly due to their short shelf life, the integration of genetic and molecular tools offers an opportunity to introgress these valuable adaptive traits into modern melon cultivars. Such efforts could lead to the development of early-season varieties with improved tolerance to environmental variability and enhanced suitability for sustainable cultivation systems. Declarations Conflict of interest statement The authors declare no conflicts of interest. Funding Declaration This research received no external funding. Author Contribution AC- design the experiment and wrote the manuscript; EO –statistical analysis; FB – vegetative material growth and handling; EA – conduct the plant pulling, ES - vegetative material growth and handling; EV- soil characterization; RNL – experimental design. Acknowledgement The authors wish to thank to the Or Emma Shemer, Yaara Sadeh, and Elad fein for help at various stages of the project. Data Availability Data will be made available on reasonable request. References Aharon, S. et al. Genetic improvement of wheat early vigor promote weed-competitiveness under Mediterranean climate. Plant Science , 303 . (2021). https://doi.org/10.1016/j.plantsci.2020.110785 Asaf, E. et al. The finger weeder cultivator for intra-row mechanical weed control: Effects of uprooting force on selected weed species. 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6","display":"","copyAsset":false,"role":"figure","size":476224,"visible":true,"origin":"","legend":"\u003cp\u003eSee image above for figure legend\u003c/p\u003e","description":"","filename":"floatimage6.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8126587/v1/d65481e08fa95ad45be0d060.jpeg"},{"id":97701242,"identity":"5cbed942-5734-419f-9558-da568b83a769","added_by":"auto","created_at":"2025-12-08 12:23:41","extension":"jpeg","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":701721,"visible":true,"origin":"","legend":"\u003cp\u003eSee image above for figure legend\u003c/p\u003e","description":"","filename":"floatimage7.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8126587/v1/858a5aaaf70777737a76fa45.jpeg"},{"id":97894494,"identity":"f5ec9bc0-981e-4483-8437-f656ddc52ef9","added_by":"auto","created_at":"2025-12-10 15:32:37","extension":"jpeg","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":533012,"visible":true,"origin":"","legend":"\u003cp\u003eSee image above for figure legend\u003c/p\u003e","description":"","filename":"floatimage8.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8126587/v1/033ed659f30de3424135bea8.jpeg"},{"id":97902520,"identity":"158b2f86-9505-4d45-b7fb-d039a642ee03","added_by":"auto","created_at":"2025-12-10 15:52:38","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":4923075,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8126587/v1/768eb04a-bdfd-48b4-b0bf-c601582795b3.pdf"},{"id":97701256,"identity":"d1d02726-f5c5-42a8-a996-15371d64f12f","added_by":"auto","created_at":"2025-12-08 12:23:41","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"supplement","size":864241,"visible":true,"origin":"","legend":"","description":"","filename":"Supplementary3.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8126587/v1/efa1ad4be3076abf1c92c9aa.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Temperature-Driven Variability in Melon Root System Architecture","fulltext":[{"header":"Introduction","content":"\u003cp\u003eIn response to climate change, many melon (\u003cem\u003eCucumis melo\u003c/em\u003e L.) growing regions have shifted sowing from the traditional summer season to late winter or early spring to reduce the adverse effects of elevated temperatures and the associated increase in evapotranspiration and pest pressure. \u003csup\u003e1,2\u003c/sup\u003e. However, early-season planting under suboptimal temperatures slows crop development, and therefore promotes weed dominance, and complicates mechanical weeding \u003csup\u003e3\u003c/sup\u003e. These challenges highlight the need to identify melon accessions with rapid early-stage growth and adaptation to suboptimal temperatures\u003c/p\u003e\u003cp\u003eMelon is one of the most popular fruits globally (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://faostat3.fao.org/\u003c/span\u003e\u003cspan address=\"http://faostat3.fao.org/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e). Its origin traces back to warm regions in Central and Southern Africa, with related species found in other warm areas, including the Indian peninsula, Asia, and Australia \u003csup\u003e4,5\u003c/sup\u003e. Consequently, melon domestication occurred independently in multiple regions \u003csup\u003e5\u003c/sup\u003e, leading to a diverse range of plant traits \u003csup\u003e6,7\u003c/sup\u003e. This variety provides a valuable resource for improving melon adaptation to different agro-technical and climatic growth conditions.\u003c/p\u003e\u003cp\u003eWeed control is a major challenge in open-field melon cultivation \u003csup\u003e8\u003c/sup\u003e. Melon is a weak competitor against weeds and experiences significant yield losses under high weed infestation levels \u003csup\u003e9\u003c/sup\u003e. Purple nutsedge (Cyperus rotundus), Puncturevine (\u003cem\u003eTribulus terrestris\u003c/em\u003e L.), Johnsongrass (\u003cem\u003eSorghum halepense\u003c/em\u003e), and pigweeds (\u003cem\u003eAmaranthus\u003c/em\u003e spp.), are just some of the summer weeds that negatively affect the growth and yield of melon. The melon\u0026rsquo;s horizontal growth habit, characterized by long, trailing vines, often fails to achieve full canopy closure through the growing season, leaving the soil exposed to light. This exposure promotes multiple weed emergence flushes and reduces melon\u0026rsquo;s ability to suppress weed growth \u003csup\u003e9\u003c/sup\u003e. Furthermore, as with many minor crops, registered herbicide options for melon are limited, providing only partial weed control. Thus, mechanical weeding remains the most effective non-chemical mean for weed removal in melon fields \u003csup\u003e8,10\u003c/sup\u003e. For mechanical weeding to be effective and safe for the crop, melon plants must have stronger resistant to pulling by the mechanical device than competing weeds.\u003c/p\u003e\u003cp\u003eCrop plants with early vigor have a competitive advantage against weeds \u003csup\u003e11\u003c/sup\u003e. Early and rapid canopy establishment suppresses weed growth by limiting their access to light \u003csup\u003e12\u003c/sup\u003e. In addition to the above-ground suppression, a well-developed root system can hinder weed emergence and establishment \u003csup\u003e13\u003c/sup\u003e. Strong root development also enhances plant resilience to environmental fluctuations, resource limitations, and various biotic and abiotic stresses \u003csup\u003e14\u0026ndash;17\u003c/sup\u003e.\u003c/p\u003e\u003cp\u003eMost early vigor studies determining differences among various varieties and/or accessions focused on wheat (\u003cem\u003eTriticum\u003c/em\u003e sp.) and other cereal crops (Aharon et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), while research on dicot crops remains limited \u003csup\u003e18\u003c/sup\u003e. To our knowledge, no previous study has examined the diversity of melon accessions for their early vigor under varying temperatures.\u003c/p\u003e\u003cp\u003eEarly vigor is strongly affected by environmental factors, particularly temperature, which drives genotype-by-environment (G \u0026times; E) interactions. Low and suboptimal temperatures are well-documented to inhibit physiological, enzymatic, and molecular processes in plants \u003csup\u003e19\u0026ndash;21\u003c/sup\u003e. Thus, identifying melon accessions with consistent root development under moderate and suboptimal temperatures has multiple advantages. It can enhance early vigor and weed competitiveness across varying climates while also ensuring safe and effective mechanical weed control. In turn, this approach could contribute to more sustainable weed management strategies in melon cultivation.\u003c/p\u003e\u003cp\u003eRoot system phenotyping is challenging due to its belowground development and the destructive nature of most measurement methods \u003csup\u003e22,23\u003c/sup\u003e. Therefore, field-based root assessments and screening breeding populations for root traits require intensive labor. In this context, root pulling using a digital force gauge provides a simple and effective first line method for assessing root system pulling strength under both field and laboratory conditions \u003csup\u003e10\u003c/sup\u003e. Given the link between pulling strength and root system structure, this method serves as a useful proxy for root system dynamics under varying conditions \u003csup\u003e24\u0026ndash;26\u003c/sup\u003e. The force required to uproot a plant is strongly correlated with key root traits such as length, diameter, biomass and other parameters \u003csup\u003e27\u0026ndash;29\u003c/sup\u003e. However, this method should be used cautiously to minimize biases introduced by soil properties (e.g., texture, soil water content) and pulling direction \u003csup\u003e29\u003c/sup\u003e.\u003c/p\u003e\u003cp\u003eSuboptimal temperatures significantly inhibit plant physiological activity. Previous studies have identified the minimum, optimum, and maximum temperatures for melon growth as 10\u0026deg;C, 34\u0026deg;C, and 45\u0026deg;C, respectively \u003csup\u003e30\u003c/sup\u003e. These conditions reduce chlorophyll production, decrease membrane fluidity, and restrict nutrient translocation, ultimately limiting plant function\u003csup\u003e20,21,31\u003c/sup\u003e. Warm-region crops, such as melons, are sensitive to moderate temperature reductions and brief frost events \u003csup\u003e32\u003c/sup\u003e. During abiotic stress, such as unfavorable temperatures, the translocation of carbohydrates and minerals from roots to shoots can \u003csup\u003e33\u0026ndash;36\u003c/sup\u003e. Therefore, physiological measurements can provide insights into differential responses among accessions, aiding in the identification of those with adaptive traits.\u003c/p\u003e\u003cp\u003eThe combined impact of suboptimal temperatures and weed competition can drastically reduce crop productivity \u003csup\u003e37\u003c/sup\u003e. While weed competitiveness remains high across temperature ranges, crop development, especially in warm-region species like melon, is significantly impaired under suboptimal temperatures. At early growth stages, the rapid expansion of weeds alters the red:far-red (R:Fr) light ratio, suppressing both above- and below-ground development of the crop. This effect can persist long after weeds are removed, leading to lasting yield reductions \u003csup\u003e38\u003c/sup\u003e. Consequently, effective and timely weed control under suboptimal temperatures is critical to preventing long-term damage and ensuring stable crop yields.\u003c/p\u003e\u003cp\u003eCurrent knowledge of melon root system diversity and function remains limited, and the relationship between root system architecture and mechanical pulling strength is still poorly understood. Only a few studies have examined variation among melon accessions in these traits. To develop sustainable strategies for mechanical weed management during early melon growth, a deeper understanding of root system characteristics and their mechanical properties is therefore essential.\u003c/p\u003e\u003cp\u003eThis study aims to screen and characterize a diverse melon population for early development, with a particular focus on performance under suboptimal temperature conditions. By integrating the pulling method, root system screening, and leaf-level gas exchange and chlorophyll fluorescence measurements, the study seeks to identify melon accessions combining tolerance to suboptimal temperatures with strong and resilient root systems.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003ePlant material and growing conditions\u003c/h2\u003e\u003cp\u003eFrom the Newe Ya'ar collection of 177 melon (\u003cem\u003eCucumis melo\u003c/em\u003e L.) accessions (Gur et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), 10 accessions representing the main variety types were selected (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Plants were grown in 2-liter pots (Pot dimensions: top diameter\u0026thinsp;=\u0026thinsp;16 cm, base diameter\u0026thinsp;=\u0026thinsp;14 cm, height\u0026thinsp;=\u0026thinsp;14 cm; volume\u0026thinsp;\u0026asymp;\u0026thinsp;2.46 l) filled with local soil (chromic Haploxerert, fine-clayey, montmorillonitic, thermic) comprising 55% clay, 25% silt, and 20% sand. Soil porosity was found to be ~\u0026thinsp;60%, the bulk density of the soil was 1.02 g cm \u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e, where field capacity and wilting point is 37% and 22% respectively (see appendix 1). The final mixture contained 2% organic matter and a pH of 7.2 (see appendix 1). Each accession was grown in five biological replicates.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eThe ten representative melon accessions used in this work from the Newe Ya\u0026rsquo;ar collection. The accessions are not subject to patent protection.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"19\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" 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namest=\"c1\"\u003e\u003cp\u003eLine Name\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e\u003cp\u003eVariety Name\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c9\" namest=\"c8\"\u003e\u003cp\u003eSub-Species\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003evarietyType\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c14\" namest=\"c13\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c16\" namest=\"c15\"\u003e\u003cp\u003eMarketing Type\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"3\" nameend=\"c19\" namest=\"c17\"\u003e\u003cp\u003eOrigin (Company)\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e\u003cp\u003eAY\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e\u003cp\u003eAnanas Yoqne'am\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c9\" namest=\"c8\"\u003e\u003cp\u003e\u003cem\u003emelo\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003eReticulatus\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c14\" namest=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c16\" namest=\"c15\"\u003e\u003cp\u003eAnanas\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c19\" namest=\"c17\"\u003e\u003cp\u003eIsrael (H\u0026rsquo;azera)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e\u003cp\u003eDUL\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e\u003cp\u003eDulce\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c9\" namest=\"c8\"\u003e\u003cp\u003e\u003cem\u003emelo\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003eReticulatus\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c14\" namest=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c16\" namest=\"c15\"\u003e\u003cp\u003eAmerican cantaloupe\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c19\" namest=\"c17\"\u003e\u003cp\u003eUSA (Bonanza)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e\u003cp\u003eNA\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e\u003cp\u003eNoy Amid\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c9\" namest=\"c8\"\u003e\u003cp\u003e\u003cem\u003emelo\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003eInodorus\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c14\" namest=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c16\" namest=\"c15\"\u003e\u003cp\u003eYellow Canary\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c19\" namest=\"c17\"\u003e\u003cp\u003eIsrael (Newe Ya\u0026rsquo;ar)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e\u003cp\u003eNY\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e\u003cp\u003eNoy Yizre'el\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c9\" namest=\"c8\"\u003e\u003cp\u003e\u003cem\u003emelo\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003eCantalupensis\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c14\" namest=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c16\" namest=\"c15\"\u003e\u003cp\u003eHa\u0026rsquo;Ogen\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c19\" namest=\"c17\"\u003e\u003cp\u003eIsrael (Newe Ya\u0026rsquo;ar)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e\u003cp\u003ePI414723\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e\u003cp\u003ePI414723\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c9\" namest=\"c8\"\u003e\u003cp\u003e\u003cem\u003eagrestis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003eMomordica\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c14\" namest=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c16\" namest=\"c15\"\u003e\u003cp\u003eN.D\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c19\" namest=\"c17\"\u003e\u003cp\u003eIndia (Pitrat M. - INRA)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e\u003cp\u003ePSR\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e\u003cp\u003ePiel De Sapo Redon\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c9\" namest=\"c8\"\u003e\u003cp\u003e\u003cem\u003emelo\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003eInodorus\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c14\" namest=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c16\" namest=\"c15\"\u003e\u003cp\u003ePiel De Sapo\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c19\" namest=\"c17\"\u003e\u003cp\u003eSpain (DEFO)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e\u003cp\u003eQME\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e\u003cp\u003eQishu Meshulash\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c9\" namest=\"c8\"\u003e\u003cp\u003e\u003cem\u003eC. callosus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003eN.D\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c14\" namest=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c16\" namest=\"c15\"\u003e\u003cp\u003eN.D\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c19\" namest=\"c17\"\u003e\u003cp\u003eIsrael (Wild accession)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e\u003cp\u003eSAS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e\u003cp\u003eSakata Sweet (17\u0026ndash;725)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c9\" namest=\"c8\"\u003e\u003cp\u003e\u003cem\u003eagrestis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003eMakuwa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c14\" namest=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c16\" namest=\"c15\"\u003e\u003cp\u003eN.D\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c19\" namest=\"c17\"\u003e\u003cp\u003eJapan (Skata)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e\u003cp\u003eTAD\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e\u003cp\u003eTAM DEW\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c9\" namest=\"c8\"\u003e\u003cp\u003e\u003cem\u003emelo\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003eInodorus\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c14\" namest=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c16\" namest=\"c15\"\u003e\u003cp\u003eHoney Dew\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c19\" namest=\"c17\"\u003e\u003cp\u003eUSA (Texas A\u0026amp;M)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e\u003cp\u003eVEP\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e\u003cp\u003eVedrantais\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c9\" namest=\"c8\"\u003e\u003cp\u003e\u003cem\u003emelo\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u003cp\u003eCantalupensis\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c14\" namest=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c16\" namest=\"c15\"\u003e\u003cp\u003eCharentais\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c19\" namest=\"c17\"\u003e\u003cp\u003eFrance (Pitrat M. - INRA)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"16\" nameend=\"c16\" namest=\"c1\"\u003e\u003cp\u003eN.D. - Not Defined\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c19\" namest=\"c17\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"17\" nameend=\"c17\" namest=\"c1\"\u003e\u003cp\u003eTable\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e. Linear correlation coefficients between days after planting and plant anchoring strength under two temperature regimes.\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c19\" namest=\"c18\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c3\" namest=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"7\" nameend=\"c10\" namest=\"c4\"\u003e\u003cp\u003e25\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"5\" nameend=\"c17\" namest=\"c13\"\u003e\u003cp\u003e16\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c19\" namest=\"c18\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c3\" namest=\"c1\"\u003e\u003cp\u003e\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003eAccession\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eSlope\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c7\" namest=\"c5\"\u003e\u003cp\u003eIntercept\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c10\" namest=\"c8\"\u003e\u003cp\u003er\u003csup\u003e2\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c14\" namest=\"c13\"\u003e\u003cp\u003eSlope\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c16\" namest=\"c15\"\u003e\u003cp\u003eIntercept\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c17\"\u003e\u003cp\u003er\u003csup\u003e2\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c19\" namest=\"c18\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c3\" namest=\"c1\"\u003e\u003cp\u003eAY\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c7\" namest=\"c5\"\u003e\u003cp\u003e2.82-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c10\" namest=\"c8\"\u003e\u003cp\u003e0.96\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c14\" namest=\"c13\"\u003e\u003cp\u003e0.53\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c16\" namest=\"c15\"\u003e\u003cp\u003e4.4-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c17\"\u003e\u003cp\u003e0.95\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c19\" namest=\"c18\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c3\" namest=\"c1\"\u003e\u003cp\u003eNY\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.72\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c7\" namest=\"c5\"\u003e\u003cp\u003e7.70-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c10\" namest=\"c8\"\u003e\u003cp\u003e0.96\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c14\" namest=\"c13\"\u003e\u003cp\u003e0.71\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c16\" namest=\"c15\"\u003e\u003cp\u003e15.66-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c17\"\u003e\u003cp\u003e0.99\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c19\" namest=\"c18\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c3\" namest=\"c1\"\u003e\u003cp\u003eDulce\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.68\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c7\" namest=\"c5\"\u003e\u003cp\u003e6.72-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c10\" namest=\"c8\"\u003e\u003cp\u003e0.86\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c14\" namest=\"c13\"\u003e\u003cp\u003e0.44\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c16\" namest=\"c15\"\u003e\u003cp\u003e8.4-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c17\"\u003e\u003cp\u003e0.92\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c19\" namest=\"c18\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c3\" namest=\"c1\"\u003e\u003cp\u003ePI414723\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.79\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c7\" namest=\"c5\"\u003e\u003cp\u003e11.08-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c10\" namest=\"c8\"\u003e\u003cp\u003e1.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c12\" namest=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c14\" namest=\"c13\"\u003e\u003cp\u003e0.12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c16\" namest=\"c15\"\u003e\u003cp\u003e1.12-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c17\"\u003e\u003cp\u003e0.98\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c19\" namest=\"c18\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003ePlants pulling\u003c/h3\u003e\n\u003cp\u003eIn the first experiment, the 10 accessions from the Newe Ya'ar collection were grown under greenhouse conditions during the winter of 2022 (planting date - January 3rd ) under natural light conditions. Temperature was set to a minimum of 18\u0026ordm;C (heating activated) and a maximum of 30 \u0026ordm;C (fan activated) with an average temperature of ~\u0026thinsp;20 \u0026ordm;C. Plants were pulled at three phenological stages: two to three true leaves, three to four true leaves, and four to five true leaves. Each measurement point (stage by accession) had five replicates.\u003c/p\u003e\u003cp\u003ePulling strength was measured using a digital force gauge (AXIS, FB500N, range 0.1\u0026ndash;50 N). The plant shoots were attached to the device with a plastic clamp and pulled horizontally, recording the maximal pulling strength for each plant. To ensure uniform soil properties, all plants were irrigated to field capacity on the morning of the pulling day, and pulled after two hours when water level is still under field capacity (figure \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e). Soil water content was measured using a true time-domain reflectometer (TDR; Acclima Inc.) following a two-point calibration (0 and 100%) conducted weekly. After the third pulling event, the soil was washed gently with water, and the root system was scanned and analyzed using WinRhizo (Regent Instruments Inc., Ottawa, ON, Canada). Roots were scan in high resulotion of 600 DPI, to ensure the detection of small diameter roots. Roots and the shoots were then oven-dried for 48 hours at 72\u0026ordm;C before weighing. Then, the specific root length (SRL, g cm\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) and specific root surface area (SRA, g cm\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e) were calculated from the WinRhizo scanning and dry weight measurement.\u003c/p\u003e\u003cp\u003eFour accessions different in their pulling strengths were selected to represent the variety in the collection. These accessions were grown in controlled environment rooms under two temperature regimes: low (16 \u0026ordm;C) and ambient (25 \u0026ordm;C). Plants were grown under artificial photosynthetic active radiation (PAR, 400\u0026ndash;700 nm) lights (~\u0026thinsp;1000 \u0026micro;mol m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e s\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, 12/12 hours day/night period), simulating early spring daylight conditions. Pulling was performed at three developmental stages, as outlined in the first experiment, with five replicates for each accession at each stage, totaling 120 plants. Roots and shoots were then dried and weighed as described above.\u003c/p\u003e\u003cp\u003eFrom the four selected accessions, two with distinct temperature responses were chosen for further study. These accessions were grown under identical controlled environmental conditions as described previously. At the four to five true leaf stage, physiological measurements, including CO₂ and H₂O fluxes, along with chlorophyll fluorescence, were conducted using the LI-COR 6800F system (Licor, Lincoln, NE, US). Pulling strength measurement, root scanning, and below and above biomass evaluation were conducted as described above, with five replicates per treatment.\u003c/p\u003e\n\u003ch3\u003eGas exchange and fluorescence parameters\u003c/h3\u003e\n\u003cp\u003eIn the second controlled environment experiment, gas exchange and fluorescence were measured using the Li-6800F system. Measurement conditions were set to 415 ppm CO₂, 40% relative humidity, a light intensity of 1000 \u0026micro;mol m⁻\u0026sup2; s⁻\u0026sup1;, and a temperature corresponding to the ambient room temperature. The youngest fully matured leaf was measured in all plants. Photosynthetic efficiency was assessed using the Pulse Amplitude Modulation (PAM) method, which was conducted simultaneously with flux measurements. To evaluate dark-adapted PAM parameters, measurements were taken in the early morning before the lights were turned on. The functionality of PSⅡ (\u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\frac{{F}_{v}}{{F}_{m}}\\)\u003c/span\u003e\u003c/span\u003e; Sipka et al., \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) was evaluated using the following equation:\u003c/p\u003e\u003cp\u003e\u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\frac{{F}_{v}}{{F}_{m}}=\\frac{{F}_{m}-{F}_{0}}{{F}_{m}}\\)\u003c/span\u003e\u003c/span\u003e [1]\u003c/p\u003e\u003cp\u003eWhere \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:{F}_{m}\\)\u003c/span\u003e\u003c/span\u003e and \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:{F}_{0}\\)\u003c/span\u003e\u003c/span\u003e are the maximal and minimal fluorescence measured under dark conditions. The light acclimated quantum yield of PSⅡ (Y(Ⅱ)) was calculated as:\u003c/p\u003e\u003cp\u003e\u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:Y\\left(Ⅱ\\right)=\\frac{{F}_{m}^{{\\prime\\:}}-\\:{F}_{0}^{{\\prime\\:}}}{{F}_{m}^{{\\prime\\:}}}\\)\u003c/span\u003e\u003c/span\u003e [2]\u003c/p\u003e\u003cp\u003eWhere \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:{F}_{m}^{{\\prime\\:}}\\)\u003c/span\u003e\u003c/span\u003e and \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:{F}_{0}^{{\\prime\\:}}\\)\u003c/span\u003e\u003c/span\u003e are the maximal and minimal fluorescence measured under light conditions.\u003c/p\u003e\u003cdiv id=\"Sec6\" class=\"Section2\"\u003e\u003ch2\u003eStatistical analysis\u003c/h2\u003e\u003cp\u003eData normality was assessed using the Shapiro-Wilk test, and homogeneity of variance was tested using Levene's test. Differences between accessions were analyzed using one-way analysis of variance (ANOVA), followed by Tukey-HSD with Bonferroni correction for multiple comparisons (α\u0026thinsp;\u0026lt;\u0026thinsp;0.0.05).\u003c/p\u003e\u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e\u003ch2\u003eVariation in pulling strength indicates differences in root morphology\u003c/h2\u003e\u003cp\u003eIn the initial screening of 10 representative melon accessions, we assessed early vigor and variation in the plant pulling force. No significant differences in pulling strength were observed among accessions at the first two phenological stages (two to four true leaves). However, at the third stage (four to five true leaves), distinct responses emerged (Figure S2). The AY and TAD accessions exhibited the highest anchoring strength, 11.52 and 9.92 N, respectively. In contrast, both the wild (QME) and commercial accessions (DUL) had the lowest values, 1.72 and 1.8 N, respectively (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). No significant effect of accession, subspecies, or variety was found on pulling strength.\u003c/p\u003e\u003cp\u003eIn addition to the plant pulling strength, we evaluated the root traits of each melon accession. A strong correlation was observed between root weight and volume in all plant accessions (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea). For example, root volumes of AY (high pulling strength) and QME (low pulling strength) were 4.25 and 2.38 cm\u003csup\u003e3\u003c/sup\u003e, respectively. However, all 10 accessions maintained a similar root-to-shoot ratio (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eb).\u003c/p\u003e\u003cp\u003eCorrelation analysis of all the measured components (root traits, weight, and their derivatives) with the pulling strength revealed that root weight (R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.77) and root volume (R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.83) were the strongest predictors of pulling strength, while specific root volume showed a weak correlation (R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.21; Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Moreover, all accessions exhibited a consistent relationship between pulling strength, root volume, and weight under these conditions (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eA-B).\u003c/p\u003e\u003cp\u003eT\u003cem\u003eemperature-driven variation in pulling strength and biomass across accessions\u003c/em\u003e\u003c/p\u003e\u003cp\u003eIn order to confirm the results of the pulling experiment, four accessions with different pulling strengths: the highest (AY), the lowest (Dulce), and two lines with moderate pulling strength (PI414723, and NY). The accessions were grown under two temperature regimes 16\u0026ordm; and 25\u0026ordm;. The pulling and weight results revealed significant variation in accession responses to temperature (Figs.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e\u0026ndash;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e, Tables\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e\u0026ndash;3).\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eLinear correlation coefficients between days after planting and plant shoot biomass under two temperature regimes.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"9\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"3\" nameend=\"c5\" namest=\"c3\"\u003e\u003cp\u003e25\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003e16\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAccession\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eSlope\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eIntercept\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003er\u003csup\u003e2\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eSlope\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eIntercept\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003er\u003csup\u003e2\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAY\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-0.80\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.97\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e1.31-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.98\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eNY\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1.81-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.99\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.14\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e3.36-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.87\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDulce\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.23\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4.17-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.90\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e3.26-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.83\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePI414723\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.24\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4.59-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.92\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.0011\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e-0.24\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.90\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eThe AY accession demonstrated the highest tolerance to suboptimal temperatures, reaching the two- to three true leaves stage 10 days earlier than the other accessions (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003eA, \u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003eA). Biomass accumulation rates remained stable at both 16\u0026deg;C and 25\u0026deg;C, with 0.1 and 0.09 g day\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e. Although NY emerged later than AY, it also exhibited high stability under both temperature conditions. Pulling strength rates were 0.72 N day\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e (16\u0026deg;C) and 0.71 N day\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e (25\u0026deg;C), while biomass accumulation rates were 0.13 g day\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e (16\u0026deg;C) and 0.14 g day\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e (25\u0026deg;C), even surpassing AY development rates.\u003c/p\u003e\u003cp\u003eIn contrast, Dulce and PI414723 accessions were highly susceptible to suboptimal temperatures. While under 25\u0026ordm;C, both accessions demonstrated the highest biomass accumulation rates (0.23 g day\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e for Dulce, 0.24 g day\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e for PI414723). However, under suboptimal temperatures, their performances declined significantly, with Dulce\u0026rsquo;s biomass accumulation reduced by ~\u0026thinsp;40%, while PI414723\u0026rsquo;s growth was nearly eliminated (~\u0026thinsp;100% reduction).\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003eThe root system response is related to the physiological response\u003c/h3\u003e\n\u003cp\u003eTo further investigate temperature response, we selected two accessions with different tolerances: stable (AY) and susceptible to suboptimal temperatures (PI414723). The AY accession was grown with five replicates per temperature, whereas PI414723 was grown with four replicates due to seedling collapse. Across both accessions, all growth parameters declined under suboptimal temperatures (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe difference in pulling strength between the tolerant and susceptible accessions was minor at 25\u0026ordm;C (7.6 N for AY and 11.2 N for PI414723) but became pronounced at 16\u0026ordm;C (6.1 N for AY and 2.8 N for PI414723; Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003ea). Root biomass did not significantly differ between the accessions across temperatures (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003eb). However, at 25\u0026ordm;C, AY exhibited significantly greater root length (1210 cm) and root volume (5.62 cm\u0026sup3;) compared to PI414723, which had lower values at both 25\u0026ordm;C (267 cm and 0.62 cm\u0026sup3;) and 16\u0026ordm;C (214 cm and 0.59 cm\u0026sup3;).\u003c/p\u003e\u003cp\u003ePhysiological measurements further supported these findings (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003e). Carbon assimilation rates declined in both accessions under suboptimal temperatures (under 16\u0026ordm;C). However, while AY showed a 40% reduction, PI414723 experienced a 70% reduction (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003ea). Stomatal conductance remained constant in AY, whereas PI414723 showed a 90% reduction at 16 \u0026ordm;C (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003eb).\u003c/p\u003e\u003cp\u003eLastly, PSⅡ functionality (\u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\frac{{F}_{v}}{{F}_{m}}\\)\u003c/span\u003e\u003c/span\u003e) decreased in both accessions, 25% in AY and 50% in PI414723 (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003ec). The PSⅡ quantum yield (Y(Ⅱ)) was reduced by 50% in AY and 65% in PI414723 under suboptimal conditions (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003ed).\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThe historical journey of melon (\u003cem\u003eCucumis melo\u003c/em\u003e) from its origins in the arid regions of Africa, through the Indian subcontinent, to East Asia and eventually the Western world, has greatly enhanced its internal diversity, ranging from physiological adaptations to the wide variability in fruit traits \u003csup\u003e5,7,40\u003c/sup\u003e. Both natural selection and human-mediated cultivation across diverse latitudes have contributed to this variability, increasing the likelihood of identifying accessions within the species that possess tolerance to both heat and cold stress \u003csup\u003e41\u003c/sup\u003e. The results of the present study highlight that even in a warm-season crop such as melon, accessions exhibiting tolerance to suboptimal temperatures can be found, likely as a result of indirect selection. The AY accession, for instance, was originally cultivated on specially prepared fallow fields under dryland conditions without irrigation, sown in early spring, and relied entirely on soil moisture conserved from winter precipitation \u003csup\u003e42\u003c/sup\u003e. This early sowing during late winter or early spring suggests that this accession possesses physiological mechanisms that enable tolerance to such unfavorable environmental conditions. Moreover, its ability to rely on residual soil moisture from winter precipitation implies the presence of a strong and efficient root system, capable of sustaining growth under limited water availability.\u003c/p\u003e\u003cp\u003eStomata closure is one of the first line process in response to temperature decrease \u003csup\u003e21,43\u003c/sup\u003e. This in turn, led to a reduction on the photosynthetic activity and decrease in the carbon assimilation rate \u003csup\u003e31,44\u003c/sup\u003e. Under optimal conditions, both accessions exhibited similar physiological behavior. However, under suboptimal conditions, their responses were distinct. While AY stomata remain open under suboptimal temperatures, the PI414723 stomatal conductance decrease sharply under same conditions. This stomatal closure induced additional feedback inhibition of photosynthesis and therefore on the carbon assimilation rate \u003csup\u003e45\u003c/sup\u003e. In addition, Zait et al., (\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) demonstrated that fluorescence parameters exhibit a gradual response to stress, with \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\frac{{F}_{v}}{{F}_{m}}\\)\u003c/span\u003e\u003c/span\u003e reduction serving as a late-stage indicator, occurring at higher stress levels. At this stage, the decrease in the xanthophyll pool may mask other parameters, such as Y(II). This can explain the similar Y(II) values observed in both accessions under suboptimal temperatures, although PI414723 was under greater stress under these conditions.\u003c/p\u003e\u003cp\u003eSeveral pathways can lead to stomata closure under suboptimal conditions as abscisic acid (ABA) and calcium movement in the plant \u003csup\u003e31,43\u003c/sup\u003e. In the current work, the AY stomata seems to be inert to the temperature reduction. This accession, cultivated for more than 100 years, challenge our understanding if it was inbreed as suboptimal temperatures accession for early sowing, or it was selected for this trait over the last decades \u003csup\u003e7,42\u003c/sup\u003e. The limited response to ABA can also explain the early emerge of the AY seedlings under sub optimal temperatures.\u003c/p\u003e\u003cp\u003eTolerance to suboptimal temperatures can be expressed in both root and shoot tissues, with physiological and molecular adjustments in one organ potentially influencing the other through systemic signaling and resource allocation \u003csup\u003e47,48\u003c/sup\u003e. Such cross-organ interactions may involve hormonal and molecular regulation, metabolic adjustments, and shifts in source\u0026ndash;sink dynamics that collectively contribute to whole-plant acclimation under suboptimal temperature conditions \u003csup\u003e49,50\u003c/sup\u003e. The current study demonstrated a direct link between aboveground physiological responses and belowground development under suboptimal temperature. In the tolerant AY accession, both root and shoot growth rates remained stable across low and ambient temperature conditions. In contrast, the susceptible PI414723 accession exhibited a concurrent reduction in root and shoot development, consistent with a marked decline in key physiological parameters.\u003c/p\u003e\u003cp\u003eEarly crop vigor is a critical trait for early-season sowing, as it enhances weed suppression and promotes successful crop establishment \u003csup\u003e11\u003c/sup\u003e. Furthermore, genotype \u0026times; environment (G \u0026times; E) stability is essential to ensure consistent crop performance under the variable environmental conditions that occur throughout the growing season \u003csup\u003e32,34\u003c/sup\u003e. The importance of maintaining such stability is becoming increasingly evident in the context of climate change, where crops must perform reliably under fluctuating and often stressful conditions. This challenge is further intensified by the global shift toward reducing chemical inputs in agricultural systems, which demands varieties with inherently robust and adaptive traits \u003csup\u003e51\u003c/sup\u003e. Although the AY accession did not exhibit the fastest development under optimal conditions, it maintained a stable developmental rate across both temperature regimes. This stability, combined with its 10-day developmental advantage under suboptimal temperatures, likely contributes to its suitability as an early-season cultivar with both agronomic and economic value. Early emergence enhances crop competitiveness and management efficiency, while also providing farmers with clear economic benefits through improved resource utilization and earlier market access.\u003c/p\u003e\u003cp\u003ePrecise and highly accurate root system phenotyping methods are often complex, costly, and require specialized equipment, which limits their accessibility and scalability for large surveys. For instance, several studies have employed X-ray computed tomography (CT; Lobet \u0026amp; Draye, \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Rellan- Alvarez et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2015\u003c/span\u003e), or custom-designed growth boxes \u003csup\u003e54\u003c/sup\u003e to investigate belowground dynamics. While these approaches provide valuable insights into root architecture and function, their major limitation lies in their unsuitability for large-scale screening, which is essential for evaluating extensive plant populations such as those examined in the present study. Despite its simplicity, the pulling method employed in this study showed a strong and consistent correlation with key root traits, particularly root volume and biomass, across all melon accessions and temperature regimes. Similar relationships between pulling strength and root system characteristics, such as total root length and volume, have also been reported in previous studies \u003csup\u003e28,29\u003c/sup\u003e. Therefore, this method represents a practical and effective tool for large-scale, first-step screening of plant populations or for field validation of more advanced and resource-demanding root phenotyping approaches. It is important to recognize that the relevance of the measured root traits extends beyond mechanical weed management, as they also play a central role in nutrient and water uptake \u003csup\u003e15,55\u003c/sup\u003e. Consequently, these traits can strongly influence plant responses to nutrient limitation and water scarcity. Future studies on root system behavior could benefit from integrating this simple pulling method with more precise phenotyping approaches, thereby enabling a more comprehensive understanding of the functional and structural characteristics of root systems under varying environmental conditions.\u003c/p\u003e\u003cp\u003eIn the present study, we successfully characterized root system responses in a large melon population using the pulling method, which was validated through root system scanning. The measurement device, a simple force gauge, is affordable, widely accessible, and easy to operate without the need for specialized training. Moreover, the method is time-efficient and provides a reliable first-line assessment of key root traits. Applying this approach as an initial screening tool prior to more labor- and time-intensive analyses could substantially improve the efficiency of studies on root dynamics and physiology.\u003c/p\u003e\u003cp\u003eFuture research on these traits is supported by the availability of fully de novo assembled and annotated genomes for the melon accessions evaluated in this study, including AY, along with readily available segregating populations \u003csup\u003e7\u003c/sup\u003e. These genomic resources provide a strong foundation for investigating the genetic basis of root pulling resistance, early seedling vigor, and temperature tolerance.\u003c/p\u003e\u003cp\u003eIntegrating these physiological and genomic insights could ultimately enable the development of melon varieties or specialized rootstocks optimized for sustainable, non-chemical weed management. Such innovations would enhance melon adaptability to both current and future environmental conditions, contributing to more resilient and resource-efficient agricultural systems.\u003c/p\u003e"},{"header":"Conclusions","content":"\u003cp\u003eThe current study demonstrated that the considerable genetic diversity within this warm-climate crop results in a wide range of responses to temperature. The limited negative response to suboptimal temperatures observed in the tolerant AY accession led to earlier emergence and higher physiological activity compared to other melon accessions. This likely explains its longstanding use as an early-season cultivar. The combination of rapid emergence and vigorous root system development enables its growth without supplemental irrigation, making it well-suited for conditions that favor early-season weed competition and mechanical weeding. Although the fruits of this accession are not compatible with current market demands, mainly due to their short shelf life, the integration of genetic and molecular tools offers an opportunity to introgress these valuable adaptive traits into modern melon cultivars. Such efforts could lead to the development of early-season varieties with improved tolerance to environmental variability and enhanced suitability for sustainable cultivation systems.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003ch2\u003eConflict of interest statement\u003c/h2\u003e\u003cp\u003eThe authors declare no conflicts of interest.\u003c/p\u003e\u003c/p\u003e\u003ch2\u003eFunding\u003c/h2\u003e\u003cp\u003eDeclaration\u003c/p\u003e\u003cp\u003eThis research received no external funding.\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eAC- design the experiment and wrote the manuscript; EO \u0026ndash;statistical analysis; FB \u0026ndash; vegetative material growth and handling; EA \u0026ndash; conduct the plant pulling, ES - vegetative material growth and handling; EV- soil characterization; RNL \u0026ndash; experimental design.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e\u003cp\u003eThe authors wish to thank to the Or Emma Shemer, Yaara Sadeh, and Elad fein for help at various stages of the project.\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eData will be made available on reasonable request.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAharon, S. et al. 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Mild salinity stimulates a stress-induced morphogenic response in Arabidopsis thaliana roots. \u003cem\u003eJ. Exp. Bot.\u003c/em\u003e \u003cb\u003e61\u003c/b\u003e (1), 211\u0026ndash;224. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/jxb/erp290\u003c/span\u003e\u003cspan address=\"10.1093/jxb/erp290\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2010).\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"diffusional limitation, pulling strength, root traits, suboptimal temperatures","lastPublishedDoi":"10.21203/rs.3.rs-8126587/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8126587/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eShifting the sowing date of summer crops such as melon (\u003cem\u003eCucumis melo\u003c/em\u003e L.) to early spring presents both agronomic opportunities and challenges. Under suboptimal temperatures, slowed physiological development allows weeds to gain a competitive advantage. Enhancing early seedling vigor under cool conditions is therefore essential for successful crop establishment and effective weed suppression. In this study, ten melon accessions representing a broader collection (~\u0026thinsp;180) were evaluated for early vigor and performance under suboptimal temperatures. Pulling resistance, root system scanning, and physiological measurements were used to identify key traits associated with early root establishment. Accessions were compared under optimal (25\u0026deg;C) and suboptimal (16\u0026deg;C) temperature regimes. Pulling strength showed a strong correlation with root biomass and volume across accessions. The AY accession exhibited clear tolerance to low temperatures, maintaining high physiological activity, whereas PI414723 displayed reduced carbon assimilation due to diffusional limitations. These findings demonstrate that tolerance to suboptimal temperatures exists even within warm-climate crops such as melon and highlight the potential of AY as a valuable genetic resource for developing early-season, stress-tolerant melon varieties.\u003c/p\u003e","manuscriptTitle":"Temperature-Driven Variability in Melon Root System Architecture","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-12-08 12:23:16","doi":"10.21203/rs.3.rs-8126587/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2026-04-08T07:09:10+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-03-24T09:12:30+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"313255756111517142323175981560912423243","date":"2026-03-16T09:23:57+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"61028674852238460527343961367839216711","date":"2026-03-11T20:07:42+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"14635650353041519017313226975301661103","date":"2026-02-12T20:48:07+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-12-20T11:37:08+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"254019269058723069500310715217700419626","date":"2025-12-05T17:10:49+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"21441275318476000992553174189254075534","date":"2025-12-05T03:32:05+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-12-04T22:18:43+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-12-04T15:03:25+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"","date":"2025-11-28T13:09:08+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-11-26T14:25:32+00:00","index":"","fulltext":""},{"type":"submitted","content":"Scientific Reports","date":"2025-11-26T14:15:43+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"8bda12f3-c379-4821-8613-8d04fa2e1953","owner":[],"postedDate":"December 8th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[{"id":59174411,"name":"Biological sciences/Ecology"},{"id":59174412,"name":"Earth and environmental sciences/Ecology"},{"id":59174413,"name":"Biological sciences/Physiology"},{"id":59174414,"name":"Biological sciences/Plant sciences"}],"tags":[],"updatedAt":"2026-04-22T12:54:20+00:00","versionOfRecord":[],"versionCreatedAt":"2025-12-08 12:23:16","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8126587","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8126587","identity":"rs-8126587","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}