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Article

Effects of Exogenous Naphthylacetic Acid Application on the Graft Union Healing of Oriental Melon Scion Grafted onto Squash Rootstock and the Qualities of Grafted Seedlings

by
Hongxi Wu
1,2,3,†,
Jingwei Liu
1,2,3,†,
Xinzhuo Miao
1,2,3,
Hao Jiang
1,2,3,
Xindi Zhang
1,2,3 and
Chuanqiang Xu
1,2,3,4,*
1
College of Horticulture, Shenyang Agricultural University, Shenyang 110866, China
2
Key Laboratory of Protected Horticulture (Ministry of Education), Shenyang Agricultural University, Shenyang 110866, China
3
Modern Protected Horticultural Engineering & Technology Center, Shenyang Agricultural University, Shenyang 110866, China
4
Key Laboratory of Horticultural Equipment (Ministry of Agriculture and Rural Affairs), Shenyang Agricultural University, Shenyang 110866, China
*
Author to whom correspondence should be addressed.
†
These authors contributed equally to this work.
Horticulturae 2025, 11(7), 765; https://doi.org/10.3390/horticulturae11070765
Submission received: 29 May 2025 / Revised: 26 June 2025 / Accepted: 30 June 2025 / Published: 2 July 2025
(This article belongs to the Section Fruit Production Systems)
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Abstract

Plant hormones are critical regulators of graft union healing, yet the specific role of exogenous naphthylacetic acid (NAA) in graft union healing efficiency and grafted seedling growth remains unexplored. In this study, we investigated the effects of exogenous NAA treatment (40 mg·L−1) on graft union healing and grafted seedling quality in oriental melon scion grafted onto squash rootstock. Our results demonstrated that exogenous NAA application significantly accelerated vascular bundle reconnection, a key indicator of successful graft union formation. The exogenous NAA treatment enhanced indole-3-acetic acid (IAA) biosynthesis by upregulating key enzymes (TDC, PDC, FMO, NIT, and TAA) and gene expression (CmYUCCA10, CmCYP450, CmoCYP450, and CmoTAA1). The exogenous NAA treatment also upregulated critical graft healing-related genes (CmoWIND1, CmoWOX4, CmoCDKB1;2, CmTMO6, CmoTMO6, CmVND7, and CmoVND7). The exogenous NAA-treated seedlings exhibited better growth. These findings reveal the potential molecular and physiological mechanisms by which exogenous NAA promotes graft union healing of melon grafted onto squash. While the results highlight the potential of exogenous NAA as a grafting enhancer under controlled conditions, further field studies are also needed to validate its practical applicability in commercial production.

1. Introduction

In horticultural crop cultivation, soil-borne diseases often reduce production and quality [1,2,3]. However, grafting technology is the critical method for solving this problem and is widely applied in horticultural crop production [4,5,6]. Grafted plants successfully deal with biological [7,8] and abiotic stresses [9,10] and increase yield and quality [11,12,13]. The success of grafting on any crop is dependent on the rootstock and scion being able to heal together effectively [14,15,16]. Moreover, it is closely related to the compatibility [17,18], genotype [19,20], and internal physiological composition [21,22] between different scion and rootstock species. The reconnection of scion and rootstock is a complicated physiological process, including isolation layer formation, callus formation, and vascular bundle formation [23,24]. Therefore, the regeneration and reconnection of the vascular bundle between scion and rootstock is an essential successful sign of graft union healing [25].
It is well known that efficiently cultivating high-quality grafted seedlings is a crucial foundation for ensuring the effectiveness of grafted cultivation [26,27,28]. Studies have shown that exogenous hormone treatment can significantly improve graft union healing efficiency and the quality of grafted seedlings [29,30]. Exogenous application of gibberellin (GA) can effectively promote cell division during tissue reunion in tomato and cucumber plants [31]. Exogenous melatonin application can accelerate the healing process of oriental melon grafted onto squash by promoting lignin accumulation [29]. Studies have shown that auxin plays a crucial role in plant tissue regeneration and vascular reconnection [32], and it is also involved in regulating xylem development and cambium formation [33]. Auxin can promote the healing of graft unions by inducing callus formation [34,35]. Rootstocks with higher endogenous auxin levels in tobacco can accelerate callus formation [36]. In Arabidopsis, the auxin of the cotyledon regulates cell proliferation in vascular tissues and cortical cell expansion to promote tissue reunion [37]. Furthermore, the auxin transport inhibitor 2,3,5-triiodobenzoic acid (TIBA, an auxin transport inhibitor) suppresses the growth of vascular cells at the graft junction [38]. Although our research group previously analyzed the effects of exogenous NAA treatment on the graft union healing of oriental melon scion grafted onto squash rootstock using transcriptomic approaches, it remains unclear whether exogenous NAA regulates graft union healing by influencing endogenous IAA biosynthesis at the graft junction. Therefore, further investigation is needed to elucidate how exogenous NAA treatment affects graft union healing.
The graft union healing of plants involves multiple physiological and molecular processes at the connection site that facilitate tissue union. Recent studies have identified several key factors regulating graft union healing in plants. AtWIND plays a crucial role in wound-induced callus formation by promoting cellular dedifferentiation and proliferation in Arabidopsis [39,40,41]. SlWOX4 may influence scion–rootstock compatibility, facilitating vascular bundle reconnection during graft healing [42]. Additionally, TMO6 is linked to callus formation and vascular regeneration in graft unions [43], while VND7 directly or indirectly activates genes involved in xylem vessel differentiation [44,45]. However, it has not been reported whether exogenous NAA application can affect graft union healing by regulating the expression of these genes related to the graft union healing of oriental melon scion grafted onto squash rootstock.
Here, we applied exogenous NAA to graft unions when oriental melon scion was grafted onto squash rootstock, and we further investigated the effects of exogenous NAA treatment on graft union healing efficiency, IAA content, the related enzyme activities of IAA biosynthesis, the key gene expression of IAA biosynthesis, the expression of genes related to graft union healing, and the quality of grafted seedlings, including the growth profiles of grafted seedlings. Furthermore, we aimed to elucidate the potential mechanisms by which exogenous NAA treatment influences the graft union healing of melon scion grafted onto squash rootstock and to provide practical technical guidance for efficiently cultivating high-quality grafted melon seedlings using the splice grafting method.

2. Materials and Methods

2.1. Plant Materials and Grafting Methods

We grafted the oriental melon cultivar ‘T0948-2’ (Cucumis melo var. Makuwa Makino) onto the squash cultivar ‘ShengZhen No.1’ (C. moschata) using the splice graft method. Grafting was performed when the scion had fully expanded its first true leaf and the rootstock had reached the cotyledon development stage, following the one-cotyledon technique [46]. During grafting, the scions were either dipped in an NAA solution (40 mg·L−1) (NAA treatment) or distilled water (control). After grafting, the seedlings were transplanted into nutrient pots (12 cm × 12 cm) and placed in a healing chamber (24 ± 2 °C, 70–80% RH, 12 h light/12 h dark, 400–600 μmol·m−2·s−1) at Shenyang Agricultural University for cultivation.

2.2. Acid Fuchsin Absorption Assay and Root Morphology Analysis

We randomly selected five plants each for analysis from the thirty plants in the NAA treatment and control groups. The stems were cross-sectioned 1.0 cm above and below the graft junction, and the rootstock stems were vertically placed in 1% acidic fuchsin solution for 1 h. Subsequently, a 2.5 mm stem segment above the graft union was cross-cut to assess acid fuchsin absorption. Observations and imaging were performed using a confocal laser microscope (Zeiss LSM 900, Jena, Germany) [29]. We randomly selected five plants from 30 grafted seedlings at the five-leaf stage in both the NAA treatment and control groups for root morphology analysis (WinRHIZO STD, Quebec, QC, Canada).

2.3. Determination of IAA Content and Enzyme Activities

We collected 0.5 g of tissues from the graft junction to quantify the levels of IAA and related enzyme activities of TDC, PDC, FMO, NIT, and TAA following the instructions provided in the enzyme-linked immunosorbent assay (ELISA) kit manual (Jiangsu Meimian Industrial Co., Ltd., Yancheng, China). Three biological replicates were performed under identical conditions.

2.4. Quantitative Real-Time PCR (qRT-PCR)

To analyze gene expression patterns during the graft union healing process, we performed qRT-PCR to quantify the relative expression levels of key genes involved in IAA biosynthesis (CmYUCCA4/6/8/10, CmCYP450, and CmTAA1 in oriental melon stems; CmoYUCCA4/6/8/10, CmoCYP450, and CmoTAA1 in squash stems) and graft union healing (CmWIND1, CmWOX4, CmCDKB1;2, CmTMO6, and CmVND7 in oriental melon stems; CmoWIND1, CmoWOX4, CmoCDKB1;2, CmoTMO6, and CmoVND7 in squash stems). The reactions were performed in a 20 µL system using a Bio-Rad CFX96 PCR instrument (Bio-Rad Laboratories, Hercules, CA, USA) with Pro Taq HS SYBR Green premix qPCR kit (AG) [47]. Primer sequences are detailed in S1. Three biological replicates were performed under identical conditions.

2.5. Statistical Analysis

All data were presented as mean ± SD (standard deviation) from three replicates. Statistical analyses and plots were performed using GraphPad Prism 8.0.2 and IBM SPSS Statistics 30.0 software, with significant differences between samples determined by one-way ANOVA (p < 0.05 or p < 0.01).

3. Results

3.1. The Exogenous NAA Treatment Accelerated the Graft Union Healing of the Oriental Melon Scion Grafted onto Squash Rootstock

In order to confirm the connectivity of the vascular bundle of the grafted seedlings, we conducted acid fuchsin absorption tests. At 6 DAG, we observed the acid fuchsin of the stem tissue sections of the oriental melon scion under the exogenous NAA treatment. However, it appeared in the control group at 8 DAG. Under the exogenous NAA treatment, all vascular bundle grafted seedlings connected at 10 DAG, but the control group did not fully connect at 10 DAG (Figure 1). In terms of the percentage of samples in which acidic fuchsin was observed, the percentage increased from 20% at 6 DAG to 100% at 9 DAG under the NAA treatment. In contrast, it increased from 40% at 8 DAG to 80% at 10 DAG in the control group (Figure S1). So, we suggested that the exogenous NAA treatment accelerated the graft union healing of oriental melon scion grafted onto squash rootstock.

3.2. The Exogenous NAA Treatment Increased the IAA Content and Related Enzyme Activities in the Graft Union Tissues

We detected the IAA (Indole-3-acetic acid) content and the enzyme activities related to IAA biosynthesis under the exogenous NAA treatment during graft union healing. The IAA content showed a trend of first increasing and then decreasing (Figure 2A). The IAA content of the control group reached the maximum at 5 DAG, while the exogenous NAA treatment reached the maximum at 6 DAG. However, it was apparent that the exogenous NAA treatment significantly increased the IAA content of the graft union tissues, except for 2 DAG. The trend of enzyme activities related to IAA synthesis, including TDC (Tryptophan decarboxylase), PDC (Pyruvate decarboxylase), FMO (Flavin-containing monooxygenase), NIT (Nitrilase), and TAA (Tryptophan aminotransferase), was generally consistent with the IAA content changes (Figure 2B,F). Furthermore, their activities under the exogenous NAA treatment were significantly higher than those of the control.

3.3. Analysis of the Key Gene Expression of IAA Biosynthesis During Graft Union Healing Under the Exogenous NAA Treatment

To further analyze the changes in the IAA content in graft union healing under the exogenous NAA treatment, we measured the expression of the YUCCA, CYP450, and TAA1 genes involved in IAA biosynthesis separately in the oriental melon scion and squash rootstock tissues (Figure 3). The results indicated that the expression of YUCCA family genes (CmYUCCA4, CmYUCCA6, CmYUCCA8, and CmYUCCA10), CmCYP450, and CmTAA1 first increased and then decreased during the graft union healing process. The exogenous NAA application differentially regulated YUCCA gene expression in the graft junction tissues. In the oriental melon scion tissues, the exogenous NAA treatment significantly downregulated CmYUCCA4 expression at 8 DAG while upregulating CmYUCCA6 at 5 DAG and CmYUCCA10 from 5 to 8 DAG (Figure 3A,B,D). The treatment showed no significant effect on CmYUCCA8 expression in the oriental melon scions (Figure 3C). In the squash rootstock tissues, the exogenous NAA treatment significantly reduced the expression levels of CmoYUCCA4 (5–8 DAG), CmoYUCCA6 (5 DAG and 6 DAG), and CmoYUCCA8 (6 DAG and 8 DAG) while transiently increasing CmoYUCCA10 expression at 2 DAG (Figure 3A–D). Additionally, the exogenous NAA treatment enhanced CmCYP450 expression in both the oriental melon scion and squash rootstock tissues at 5 and 8 DAG (Figure 3E) and upregulated CmTAA1 expression in the oriental melon scion tissues at 6 DAG and in the squash rootstock tissues from 2 to 6 DAG (Figure 3F).

3.4. Expression Profiles of the Genes Related to Graft Union Healing of the Oriental Melon Scion Grafted onto Squash Rootstock by the Exogenous NAA Treatment

To further analyze the effects of the exogenous NAA treatment on the graft union healing of the oriental melon scion grafted onto squash rootstock, we determined the expression of the genes related to graft union healing, including WOUND-INDUCED DEDIFFERENTIATION1 (WIND1), WUSCHEL-RELATED HOMEOBOX4 (WOX4), CYCLIN-DEPENDENT KINASE (CDKB1;2), TARGET of MONOPTEROS 6 (TMO6), and VASCULAR-RELATED NAC-DOMAIN 7 (VND7) of the oriental melon and squash rootstock tissues during graft union healing (Figure 4). The results indicated that the exogenous NAA treatment did not significantly affect the expression profiles of CmWIND1, CmWOX4, and CmCDKB1;2 (Figure 4A–C). The relative expression of CmWIND1 and CmoWIND1 gradually increased from NG to 2 DAG. The exogenous NAA treatment significantly increased the relative expression of CmoWIND1 from 1 DAG to 2 DAG (Figure 4A). The relative expression profiles of CmWOX4 and CmoWOX4 were similar to those of CmWIND1 and CmoWIND1. The exogenous NAA treatment significantly improved the relative expression of CmoWOX4 at 1 DAG and 3 DAG (Figure 4B). From 5 DAG to 6 DAG, the exogenous NAA treatment significantly increased the relative expression of CmoCDKB1;2 (Figure 4C). Under the exogenous NAA treatment, the relative expression of CmTMO6 was significantly higher than the control from 6 DAG to 7 DAG, and the relative expression of CmoTMO6 was significantly higher than the control at 6 DAG and 8 DAG, respectively. From 8 DAG to 9 DAG, the relative expression of CmVND7 and CmoVND7 was significantly higher than the control under the exogenous NAA treatment.

3.5. Effects of the Exogenous NAA Treatment on the Growth Performance of the Grafted Oriental Melon Seedlings

The quality of grafted seedlings is the basis of ensuring the quality and yield of oriental melon fruit. It is unknown whether the exogenous NAA treatment affected the grafted oriental melon seedlings, although the exogenous NAA promoted the graft union healing of the oriental melon scion grafted onto squash rootstock. As shown in Figure 5A, the exogenous NAA treatment significantly enhanced the growth vigor of the grafted oriental melon seedlings compared to the control. The exogenous NAA treatment significantly increased the fresh and dry weight of the roots, stems, and leaves of the grafted oriental melon seedlings (Figure 5B,C). We also investigated the growth status of the grafted oriental melon seedlings (Figure 6A). The results showed that the exogenous NAA treatment significantly increased the total root length (Figure 6B), the root surface (Figure 6C), the root volume (Figure 6D), the main root length (Figure 6G), and the total number of root tips and forks (Figure 6E,F). Evidently, the grafted seedlings treated with the exogenous NAA developed more robust root systems, contributing to the grafted oriental melon plants’ vigorous growth.

4. Discussion

Grafting is an indispensable technique in modern horticulture for propagating elite cultivars, enhancing stress tolerance, and improving fruit quality, particularly in economically important crops like oriental melon [48]. The success of grafting hinges on the rapid and effective healing of the graft union, a complex process involving wound response, callus formation, and vascular redifferentiation [29,49]. This study demonstrated that exogenous naphthylacetic acid (NAA) application significantly promoted the graft union healing process in oriental melon scions grafted onto squash rootstocks. Our findings revealed that NAA not only accelerated the physical healing of the graft junction but also enhanced endogenous indole-3-acetic acid (IAA) levels, modulated the activity of related enzymes, altered the expression of key genes involved in IAA biosynthesis and graft healing, and ultimately improved the overall growth performance of the grafted seedlings.
The observed acceleration of graft union healing by exogenous NAA application was consistent with the well-established role of auxins in promoting cell division, differentiation, and wound healing in plants [30]. Auxins, including NAA, stimulate callus proliferation at the wound site, a critical initial step for bridging the scion and rootstock [50,51]. Subsequently, auxins play a pivotal role in the differentiation of vascular tissues (xylem and phloem) across the callus bridge, re-establishing symplastic and apoplastic continuity essential for water and nutrient transport [52,53]. Our results aligned with previous work on oriental melon grafted onto squash, where NAA treatment was shown to hasten the healing process [30]. In plants, factors such as temperature and other growth regulators like melatonin influence this process [29,54], highlighting the complex interplay of internal and external cues. As observed in our study, the efficiency of NAA application in promoting healing underscores its potential as a practical tool in grafting technology. However, because of limitations, such as experimental conditions, it remains necessary to conduct extensive field trials and validation under various graft union healing environment conditions to determine whether exogenous NAA treatment can consistently and significantly improve graft union healing efficiency and the quality of melon grafted onto squash in practical production. This is essential to ensure its effectiveness in agricultural applications.
Delving into the molecular mechanisms, the exogenous application of NAA significantly promoted IAA biosynthesis by upregulating both key biosynthetic enzymes (TDC, PDC, FMO, NIT, and TAA) and associated genes (CmYUCCA10, CmCYP450, CmoCYP450, and CmoTAA1). Additionally, the exogenous NAA treatment enhanced the expression of genes involved in graft union healing, including CmoWIND1, CmoWOX4, CmoCDKB1;2, CmTMO6, CmoTMO6, CmVND7, and CmoVND7. This finding was crucial as it indicated that NAA acted as auxin and potentiated the plant’s intrinsic auxin production machinery at the critical site of union formation. Beyond IAA biosynthesis, NAA influenced a broader spectrum of genes associated with graft union healing. These likely included genes regulating cell cycle progression, essential for callus proliferation [29]. Additionally, genes involved in vascular redifferentiation, such as VASCULAR-RELATED NAC-DOMAIN 7 (VND7) and TARGET OF MONOPTEROS 6 (TMO6), were critical for establishing functional xylem and phloem connections [29]. Cell wall-modifying enzymes, such as cellulases and pectinases, are also vital for cell separation, expansion, and the subsequent fusion of tissues at the graft interface [48] and affect genes in plant hormone signal transduction, phenylpropanoid biosynthesis, and phenylalanine metabolism pathways [30]. Similarly, studies on pear grafting identified numerous differentially expressed genes related to cell wall metabolism, hormone signaling, and stress responses during union formation [49]. AINTEGUMENTA-like (AIL) transcription factors have also been implicated in pumpkin graft union healing, highlighting the role of specific regulatory networks [55]. The NAA-induced modulation of these gene networks orchestrates the complex cellular events leading to successful graft take. It is worth noting that genes, such as WIND1, WOX4, CDKB1;2, TMO6, and VND7, that are potentially involved in directly regulating the graft union healing of melon grafted onto squash still require further functional validation during the graft union healing process. This will enable a deeper understanding of the mechanism by which exogenous NAA treatment influences graft union healing of melon grafted onto squash.
The accelerated and improved healing of the graft union, facilitated by the NAA application, logically translated into enhanced growth performance of the grafted oriental melon seedlings. A well-formed vascular connection ensures efficient water, mineral, and photosynthate transport between the rootstock and scion, which is fundamental for vigorous growth [56]. Our findings agreed with Xu et al. [30], who reported improved growth characteristics in NAA-treated grafted melon seedlings. Producing high-quality, vigorous seedlings was a primary goal in commercial nurseries, and NAA application appeared to be a viable strategy to achieve this. This is particularly important as the initial quality of grafted seedlings can significantly impact their subsequent field establishment, stress resilience, and eventual yield. Other plant growth regulators and treatments were also explored for similar benefits, such as NAA application for rooting and shoot elongation in poplar tissue culture [57] or using NAA treatment in controlled-release formulations to promote root formation [58]. However, in our experiment, we did not measure indicators such as the photosynthetic characteristics or water use efficiency of the grafted seedlings. Further studies are needed to clarify the photosynthetic performance and other key traits of grafted melon seedlings under exogenous NAA treatment in order to better evaluate the practical application potential of NAA in cultivating high-quality grafted melon seedlings.
The exogenous NAA application offered a simple, cost-effective, and efficient method to improve grafting success rates and produce higher-quality seedlings in cucurbit production from a practical standpoint. However, this study has certain limitations. The effects of NAA are often concentration-dependent, and while our study demonstrates efficacy, further optimization of the NAA concentration, timing, and application method could yield even better results. Our assessment was primarily at the seedling stage. Long-term evaluations of field performance, including plant vigor, stress tolerance, fruit yield, and quality, are necessary to ascertain the full benefits of NAA treatment. While gene expression analysis provided molecular insights, a more comprehensive understanding could be achieved through proteomic and metabolomic profiling of the graft union during NAA-induced healing. The interaction of NAA with environmental factors, such as temperature and humidity [59,60], also needs to be considered for robust practical applications. Future research should aim to address these limitations. During graft union healing, investigating the interplay between NAA and other endogenous plant hormones, such as cytokinins, gibberellins, ethylene, and brassinosteroids, could uncover more complex regulatory networks [48]. The role of sugar signaling and metabolism, which is increasingly recognized as crucial for graft healing [48,54], in conjunction with NAA action, presents another exciting avenue. Functional characterization of the identified NAA-responsive genes using genetic approaches (like CRISPR/Cas9-mediated gene editing or overexpression/silencing lines) will be essential to confirm their specific roles in the grafting process. Furthermore, exploring novel delivery systems for NAA, such as nano-formulations [58], could enhance its efficacy and sustainability in horticultural practice.

5. Conclusions

This study demonstrated that exogenous NAA application can enhance graft union healing and improve early seedling development in oriental melon grafted onto squash rootstock under controlled conditions. The observed improvements were associated with modulated endogenous IAA levels, enzymatic activities, and the regulation of key genes involved in cell proliferation, differentiation, and vascular regeneration. However, these findings were based on short-term experiments, and further validation under field conditions is necessary to assess the long-term effects, cost-effectiveness, and practical applicability of NAA treatment in commercial grafting. While these results provided valuable insights into the hormonal and molecular mechanisms underlying graft healing, their broader horticultural implications remained to be evaluated.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/horticulturae11070765/s1, Table S1: Primers sequences of qRT-PCR. Figure S1: The percentage of acid fuchsin detected during graft union healing of oriental melon scion grafted onto squash rootstock.

Author Contributions

H.W. and J.L.: investigation, data curation, validation, software, writing—original draft; X.M., H.J. and X.Z.: investigation; C.X.: project administration, supervision, writing—review and editing. All authors have read and agreed to the published version of the manuscript.

Funding

This work was supported by the National Natural Science Foundation of China (32272696) and the Department of Science & Technology of Liaoning province (2023JH1/10200010).

Data Availability Statement

The data presented in this study are available on request from the corresponding author.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. Observation of acid fuchsin absorption during the graft union healing of oriental melon scion grafted onto squash rootstock. NAA, naphthylacetic acid; DAG, days after grafting; scale bars, 1 mm.
Figure 1. Observation of acid fuchsin absorption during the graft union healing of oriental melon scion grafted onto squash rootstock. NAA, naphthylacetic acid; DAG, days after grafting; scale bars, 1 mm.
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Figure 2. The changes in the IAA content and related enzyme activities in the graft union tissues under the exogenous NAA treatment. (A), IAA content. (B), TDC activities. (C), PDC activities. (D), FMO activities. (E), NIT activities. (F), TAA activities. NAA, naphthylacetic acid. DAG, days after grafting. Different lowercase letters indicated significant differences, p < 0.05.
Figure 2. The changes in the IAA content and related enzyme activities in the graft union tissues under the exogenous NAA treatment. (A), IAA content. (B), TDC activities. (C), PDC activities. (D), FMO activities. (E), NIT activities. (F), TAA activities. NAA, naphthylacetic acid. DAG, days after grafting. Different lowercase letters indicated significant differences, p < 0.05.
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Figure 3. Analysis of the key gene expression of IAA biosynthesis in oriental melon scion and squash rootstock tissues during graft union healing under exogenous NAA treatment. (A) CmYUCCA4 and CmoYUCCA4. (B), CmYUCCA6 and CmoYUCCA6. (C), CmYUCCA8 and CmoYUCCA8. (D), CmYUCCA10 and CmoYUCCA10. (E), CmCYP450 and CmoCYP450. (F), CmTAA1 and CmoTAA1. NAA, naphthylacetic acid. DAG, days after grafting. Different lowercase letters indicate significant differences, p < 0.05.
Figure 3. Analysis of the key gene expression of IAA biosynthesis in oriental melon scion and squash rootstock tissues during graft union healing under exogenous NAA treatment. (A) CmYUCCA4 and CmoYUCCA4. (B), CmYUCCA6 and CmoYUCCA6. (C), CmYUCCA8 and CmoYUCCA8. (D), CmYUCCA10 and CmoYUCCA10. (E), CmCYP450 and CmoCYP450. (F), CmTAA1 and CmoTAA1. NAA, naphthylacetic acid. DAG, days after grafting. Different lowercase letters indicate significant differences, p < 0.05.
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Figure 4. The relative expression of genes related to graft union healing in the melon scions and squash rootstock during the graft healing process. (A) CmWIND1 and CmoWIND1. (B), CmWOX4 and CmoWOX4. (C), CmCDKB1;2 and CmoCDKB1;2. (D), CmTMO6 and CmoTMO6. (E), CmVND7 and CmoVND7. NAA, naphthylacetic acid. DAG, days after grafting. Different lowercase letters indicate significant differences, p < 0.05.
Figure 4. The relative expression of genes related to graft union healing in the melon scions and squash rootstock during the graft healing process. (A) CmWIND1 and CmoWIND1. (B), CmWOX4 and CmoWOX4. (C), CmCDKB1;2 and CmoCDKB1;2. (D), CmTMO6 and CmoTMO6. (E), CmVND7 and CmoVND7. NAA, naphthylacetic acid. DAG, days after grafting. Different lowercase letters indicate significant differences, p < 0.05.
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Figure 5. The changes in the fresh and dry weight of the grafted seedlings after the exogenous NAA treatment. (A), The growth performance of the grafted seedlings. (B), he fresh weight of the grafted seedlings. (C), The dry weight of grafted seedlings. NAA, naphthylacetic acid. Significant differences are indicated with * p < 0.05.
Figure 5. The changes in the fresh and dry weight of the grafted seedlings after the exogenous NAA treatment. (A), The growth performance of the grafted seedlings. (B), he fresh weight of the grafted seedlings. (C), The dry weight of grafted seedlings. NAA, naphthylacetic acid. Significant differences are indicated with * p < 0.05.
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Figure 6. Analysis of the root growth status of the grafted seedlings after the exogenous NAA treatment. (A), Photographs of roots. (B), The total root length. (C), The total root surface area. (D), The total root volume. (E), The total number of root tips. (F), The total number of root forks. (G), The main root length. NAA, naphthylacetic acid. Significant differences are indicated with * p < 0.05 and ** p < 0.01.
Figure 6. Analysis of the root growth status of the grafted seedlings after the exogenous NAA treatment. (A), Photographs of roots. (B), The total root length. (C), The total root surface area. (D), The total root volume. (E), The total number of root tips. (F), The total number of root forks. (G), The main root length. NAA, naphthylacetic acid. Significant differences are indicated with * p < 0.05 and ** p < 0.01.
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Wu, H.; Liu, J.; Miao, X.; Jiang, H.; Zhang, X.; Xu, C. Effects of Exogenous Naphthylacetic Acid Application on the Graft Union Healing of Oriental Melon Scion Grafted onto Squash Rootstock and the Qualities of Grafted Seedlings. Horticulturae 2025, 11, 765. https://doi.org/10.3390/horticulturae11070765

AMA Style

Wu H, Liu J, Miao X, Jiang H, Zhang X, Xu C. Effects of Exogenous Naphthylacetic Acid Application on the Graft Union Healing of Oriental Melon Scion Grafted onto Squash Rootstock and the Qualities of Grafted Seedlings. Horticulturae. 2025; 11(7):765. https://doi.org/10.3390/horticulturae11070765

Chicago/Turabian Style

Wu, Hongxi, Jingwei Liu, Xinzhuo Miao, Hao Jiang, Xindi Zhang, and Chuanqiang Xu. 2025. "Effects of Exogenous Naphthylacetic Acid Application on the Graft Union Healing of Oriental Melon Scion Grafted onto Squash Rootstock and the Qualities of Grafted Seedlings" Horticulturae 11, no. 7: 765. https://doi.org/10.3390/horticulturae11070765

APA Style

Wu, H., Liu, J., Miao, X., Jiang, H., Zhang, X., & Xu, C. (2025). Effects of Exogenous Naphthylacetic Acid Application on the Graft Union Healing of Oriental Melon Scion Grafted onto Squash Rootstock and the Qualities of Grafted Seedlings. Horticulturae, 11(7), 765. https://doi.org/10.3390/horticulturae11070765

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