1 Introduction

Changshan Huyou (Citrus changshan-huyou) is a hybrid citrus fruit tree unique to Zhejiang Province. It has a unique flavor and is rich in active ingredients. It has attracted attention due to its strong adaptability and high medicinal value (Gao et al., 2022). The fruit of Changshan Huyou can not only be eaten fresh and processed, but also be widely used in the food, medicine and health care products industries because it contains components such as naringin, naringin and limonin (Wang et al., 2024). This kind of fruit tree originated in Changshan County, Zhejiang Province. It has a cultivation history of over a hundred years. Now it has become a local characteristic industry and has been listed as a geographical indication protected product of China (Gao et al., 2022; Miao et al., 2024). With the expansion of the planting scale, the cultivation area of Changshan Huyou has gradually expanded to Zhejiang, Hunan, Jiangxi and other places, forming a diverse ecological adaptation pattern (Tan et al., 2012; Yu et al., 2012).

Ecological conditions have a significant impact on the fruit traits and yield of Changshan Huyou. Factors such as climate, soil and terrain in different regions cause significant differences in fruit size, nutritional components and flavor substances (Huang et al., 2025; Ke et al., 2025). For instance, temperature, rainfall and soil fertility not only affect fruit size, soluble solids and acidity, but also influence the accumulation of volatile components and bioactive substances (Gao et al., 2022; Wang et al., 2024). Meanwhile, the adaptation of Changshan Huyou in different ecological zones also provides references for industrial layout and variety improvement (Tan et al., 2012; Yu et al., 2012).

This study summarizes the differences in fruit traits and regional manifestations of Changshan Huyou in different ecological zones, discusses the influence mechanism of ecological conditions on fruit quality and yield, and points out that these research results provide a scientific basis for the development of the industry. This study integrates the latest achievements in molecular omics, morphology and ecology, aiming to provide theoretical support for the variety selection and breeding, regional optimization and high-quality industrial development of Changshan Huyou.

2 Biological Characteristics and Fruit Traits of Changshan Huyou

2.1 Morphological features: tree architecture, flowering, and fruiting patterns

Changshan Huyou is an evergreen subtropical fruit tree with a strong body, a large crown and vigorous growth of branches. The leaves are thick and elliptical, with strong adaptability, cold resistance and good disease and pest resistance (Zhong, 2004; Tan et al., 2012). Flowering mainly occurs in spring, with large floral organs, high pollen vitality, normal development of stamens and pistils, and relatively high pollination and fertilization rates. The flowers and fruits of the natural tetraploid Changshan Huyou are larger than those of ordinary plants, and the pollen grains are larger but slightly less vigorous. There is little difference in the development of the pistil, which indicates that it is of great value in breeding and hybridization (Huang et al., 2025). The fruit’s expansion period mainly occurs from June to July and September, and it ripens from October to November. The single fruit has a large weight, thick skin, and is resistant to storage and transportation (Gao et al., 2022; Miao et al., 2024).

2.2 Key fruit quality traits: size, weight, peel thickness, juice content, sugar-acid balance, and aroma

The fruit of Changshan Huyou is pear-shaped, spherical or flat-spherical, with a single fruit weighing about 300 grams. Tetraploid fruits can reach twice the size of diploid ones (Chen et al., 2011; Huang et al., 2025). The peel is relatively thick, which is conducive to storage and transportation, but it will affect the consumption rate (Kang et al., 2022). The fruit flesh is juicy, the juice sacs are plump, the sugar-acid ratio is moderate, and the taste is sweet, sour and slightly bitter. It also has the characteristic aroma of pomelo (Miao et al., 2024; Wang et al., 2024). The content of soluble solids (SSC) is generally between 10 and 13 Brix. The SSC of tetraploid fruits is slightly lower, but the difference in organic acids is not significant (Huang et al., 2025; Ke et al., 2025). The fruit is rich in volatile organic compounds (VOCs) and has diverse aroma substances, which will change during ripening and storage. Among them, limonene, naringin and hesperidin are the main components (Gao et al., 2022; Wang et al., 2024). Traits such as fruit size, skin thickness and juice amount vary among different regions and varieties (Chen et al., 2011; Tan et al., 2012).

2.3 Commercial and nutritional values of fruit traits

The fruit of Changshan Huyou has an attractive appearance, is large in size and golden in color. It is suitable for storage and transportation, and is suitable for fresh consumption as well as deep processing (Zhong, 2004; Chen et al., 2011). The peel and pulp of the fruit contain various active substances, such as naringin, hesperidin,limonin, volatile oil, flavonoids and terpenoids, etc. (Miao et al., 2024). These components have antioxidant, anti-inflammatory, hypoglycemic, liver-protecting, anti-tumor and other effects (Figure 1) (Gao et al., 2022; He et al., 2023; Huang et al., 2025). The fruits and by-products of Changshan Huyou (such as peel and residue) have been widely used in the food, medicine and health care products industries, and their economic and nutritional values are prominent (Figure 2) (Zhang et al., 2012; Gao et al., 2022). In addition, the fruit contains a considerable amount of soluble sugar, rich vitamin C and dietary fiber, which further enhances its nutritional quality (Li et al., 2020b; Miao et al., 2024).

3 Ecological Zoning of Changshan Huyou Cultivation

3.1 Major ecological zones: climate, soil, and topography differences

Changshan Huyou is mainly distributed in Quzhou, Zhejiang Province in southeastern China and the surrounding hilly areas. This place has a subtropical monsoon climate with moderate temperature and abundant rainfall, which is very suitable for the growth of citrus fruit trees (Tan et al., 2012). The local soil is mainly red soil, with deep layers, good drainage and relatively high organic matter content. However, there is a problem of soil erosion on some slopes (Chen et al., 2004). The terrain is mostly low mountains and hills, with an altitude generally ranging from 100 to 500 meters. The slope is not steep, which is conducive to air circulation and light distribution. However, more attention should be paid to soil and water conservation when planting on sloping land. Considering the comprehensive climatic, soil and topographic conditions, Quzhou and its surrounding hills are the most suitable for growing Changshan Huyou. More than 65% of the local area has been designated as suitable or sub-suitable planting areas (Yu et al., 2012).

3.2 Historical development of regional planting practices

Changshan Huyou originated in Quzhou, Zhejiang Province, where it has a long history of cultivation. Since the 20th century, with the improvement of varieties and the advancement of cultivation techniques, the planting areas have gradually expanded. Some newly introduced regions can also achieve stable output and good quality (Tan et al., 2012). In recent years, due to climate change and land use adjustment, the planting area in the peripheral regions has fluctuated, but the scale and output of the core production areas remain stable (Yu et al., 2012). Historically, the expansion of the Changshan Huyou planting area mainly relied on conditions such as suitable climate, soil fertility and hilly terrain. High-quality orchards are often concentrated in areas with better ecological environment and better resource conditions (Chen et al., 2004).

3.3 Regional resource advantages and limitations

The main production area of Changshan Huyou has many natural advantages. Firstly, the climate is warm and humid with abundant rainfall, which is suitable for fruit enlargement and quality improvement. Secondly, red soil is rich in organic matter, which is conducive to root growth. Thirdly, hilly terrain is conducive to drainage and can also reduce pests and diseases (Yu et al., 2012). In addition, the biodiversity of the local orchard ecosystem is relatively high, which is conducive to improving stress resistance and ecological stability (Chen et al., 2004). However, there are also some problems: sloping land is prone to soil erosion and a decline in soil fertility. Extreme weather conditions (such as drought and rainstorm) can affect yield and quality (Yu et al., 2012). In addition, the climate and soil conditions in the marginal area vary greatly, and the suitability for planting is limited (Tan et al., 2012). Therefore, scientific ecological regionalization and rational utilization of resources are the keys to the sustainable development of Changshan Huyou industry.

4 Variations in Yield Performance Across Zones

4.1 Regional differences in fruit yield per hectare

Changshan Huyou is a unique citrus crop in China, and its yield performance varies significantly in different ecological zones. In its original production area of Quzhou, Zhejiang Province, Changshan Huyou often maintains high yields, good quality and strong adaptability due to the favorable climate, soil and management conditions (Tan et al., 2012). The regional evaluation results show that the area of the suitable zone exceeds 65%, the area of the sub-suitable zone is about 13%, and the area of the unsuitable zone is 21%, indicating that ecological zoning plays an important role in the distribution of yield (Yu et al., 2012). In the introduced cultivation areas, such as Chenzhou, Hunan Province, Changshan Huyou can also grow normally and maintain a stable yield, which indicates that it has a strong adaptability to the subtropical climate in the south (Tan et al., 2012). In addition, the pH and organic matter content of the soil have a significant impact on the yield. Reasonable soil management can further increase the yield per unit area (Wu et al., 2024).

4.2 Seasonal stability and inter-annual yield fluctuations

The output of Changshan Huyou is not only affected by the regional ecological conditions, but also experiences seasonal and interannual fluctuations. Climate change, precipitation and temperature will directly affect fruit enlargement, sugar-acid ratio and final yield (Dong et al., 2024). Studies have found that moderate water stress management, such as staged drip irrigation, can improve water use efficiency without reducing yield, thereby achieving water conservation and high yield (Chen et al., 2025). On an interannual scale, extreme climate events such as droughts or rainstorms can cause fluctuations in production. However, through reasonable irrigation and fertilization management, these adverse effects can be mitigated to maintain the stability and sustainability of production (Tong et al., 2022; Devin et al., 2023). Meanwhile, increasing variety diversity and adopting grafting techniques can also help stabilize yields (Reiss and Drinkwater, 2018; Ke et al., 2025).

4.3 Long-term productivity sustainability in diverse zones

The long-term productivity of Changshan Huyou depends on multiple factors such as climate, soil, management methods and genetic diversity of varieties. Long-term monitoring and simulation studies have shown that reasonable ecological zone management, such as optimizing planting layout, selecting appropriate varieties and implementing ecological compensation, is conducive to maintaining high yields while taking ecological protection into account (Yu et al., 2012; Yu and Wu, 2018; Xie et al., 2023). Measures such as ecological intensive management, crop rotation, intercropping and organic management can improve soil health, increase microbial diversity, and enhance the stress resistance and sustainability of the system (MacLaren et al., 2022; Janke et al., 2024). Meanwhile, variety improvement based on molecular breeding and genomics has also provided new means to enhance the stress resistance and long-term yield of Changshan Huyou (Miao et al., 2024). In the future, more attention needs to be paid to how to balance the sustainability of production and ecological environment protection in the context of climate change.

5 Variations in Fruit Quality and Commercial Traits

5.1 Differences in fruit size, shape, and peel characteristics among zones

The fruit morphology of Changshan Huyou varies greatly in different ecological zones. Studies have found that tetraploid fruits are larger than diploid ones, with significant increases in fruit diameter and height, rounder shape, thicker fruit skin, and the average weight is almost twice that of diploid ones (Huang et al., 2025). In some areas, the fruit skin color is brighter and more evenly distributed, enhancing the appearance and commercial value of the fruit (Ke et al., 2025). These morphological differences are not only related to genetics, but also have a great relationship with climate and soil conditions (Yu et al., 2012). Introduction studies have shown that Changshan Huyou grows well in the humid and warm southern regions, with normal fruit development, stable commercial traits, and few diseases and pests, making it suitable for large-scale promotion (Tan et al., 2012).

5.2 Regional influences on sugar-acid ratio, nutritional compounds, and flavor

The sugar-acid ratio, nutrition and flavor of Changshan Huyou also vary depending on the ecological area. Pomelo fruits grafted with Changshan Huyou as the intermediate rootstock have a higher content of soluble solids, lower acidity, better flavor, and more prominent flesh color and aroma (Ke et al., 2025). The soluble solids of tetraploid Changshan Huyou are slightly lower than those of diploid, but the content of organic acids is close. Meanwhile, some amino acids, flavonoids and alkaloids will increase significantly in specific regions and varieties (Huang et al., 2025). The fruit is also rich in naringin, hesperidin and limonin. The contents of these functional substances are affected by climate, soil and management methods, which determine the flavor and medicinal value of the fruit (Gao et al., 2022; Miao et al., 2024). Furthermore, the regional environment can also affect the composition of volatile organic compounds (VOCs) in fruits, thereby altering the complexity of aroma and flavor (Wang et al., 2024; Ke et al., 2025).

5.3 Storage ability, shelf life, and transport adaptability under different ecological conditions

The fruit of Changshan Huyou is renowned for its good storability, long shelf life and excellent transportability. Studies have shown that within 60 days of postharvest storage, the oxidative monoterpenoids in fruit peel essential oils decrease while the sesquiterpenoids increase, enhancing antioxidant and antibacterial activities. This helps to extend the shelf life and maintain quality (Wang et al., 2024). The peel is thick and the structure is compact, which can also reduce damage and rot during transportation (Huang et al., 2025). In suitable ecological areas, Changshan Huyou has better storability, a higher commercial rate, and is suitable for long-distance transportation and market sales (Tan et al., 2012; Yu et al., 2012). In addition, the maturity of the fruit, the time of harvest and the storage environment will all affect its storability and shelf life (Gao et al., 2022; Wang et al., 2024).

6 Environmental and Management Drivers of Variation

6.1 Climatic factors: temperature, rainfall, and sunlight effects on fruit traits

Citrus fruits show significant differences in different ecological regions. Climatic factors are one of the main reasons for the changes in the fruit traits of Changshan Huyou. Temperature directly affects the growth cycle and ripening speed of fruits. At high temperatures, the fruits ripen faster, but the organic acids decrease, the taste becomes sweeter, the storage period of the fruits shortens, and they are more prone to rot or uneven color. Rainfall and water supply can affect fruit size and sugar accumulation. When there is sufficient water, the fruit is large and juicy. However, if there is too much rain, the sugar will be diluted and the flavor will become weak (Sadka et al., 2019). Light is also very important. An appropriate amount of light can make the fruit color better and the sugar content higher, but too strong light or insufficient light will affect the quality (Feng et al., 2021). In addition, climate change (such as warming in winter and increased extreme weather) also has a negative impact on the quality of citrus fruits, which requires special attention (Sadka et al., 2019; Zhang et al., 2022).

6.2 Soil fertility, pH, and water management impacts

Soil conditions are equally crucial. Soils with high fertility and rich minerals (such as potassium, iron, copper, etc.) can make fruits more nutritious and have better flavors. The pH value of the soil is also very important. Slightly acidic to neutral is most suitable for citrus growth (Czech et al., 2019). Water management should be balanced. It is necessary to ensure fruit enlargement and sugar accumulation, while avoiding excessive watering that causes root hypoxia and thus affects quality (Figure 3) (Sadka et al., 2019). Due to the different soil conditions and water management methods in different ecological zones, the fruit traits also show significant differences (Zhang et al., 2022).

6.3 Cultivation practices: pruning, fertilization, and pest control in different zones

The influence of cultivation and management measures on fruit traits is becoming increasingly significant. Scientific pruning can improve the tree shape, increase ventilation and light, and make the fruits more uniform and of better quality (Feng et al., 2021). Rational fertilization, especially the balanced supply of nitrogen, phosphorus and potassium, will directly affect the size, sugar-acid ratio and nutritional components of the fruit (Czech et al., 2019). Pest and disease control is also very important, as it determines the health of the fruit and the commercial rate. Nowadays, many orchards have begun to use precise pesticide application or biological control to reduce pesticide residues and make fruits safer (Dhiman et al., 2023). Due to the differences in climate, soil and types of pests and diseases in different regions, the cultivation methods need to be adapted to local conditions to form a technical system suitable for the local area (Zhang et al., 2022).

7 Case Study: Regional Performance Evaluation of Changshan Huyou

7.1 Study background: site selection, ecological characteristics, and management systems

Changshan Huyou is a unique citrus fruit tree in Zhejiang Province, with strong ecological adaptability and both medicinal and economic values (Gao et al., 2022; Miao et al., 2024; Huang et al., 2025). The main production area is in Changshan County, Quzhou City, Zhejiang Province and its surrounding areas. Later, it was also introduced to Hunan, Jiangxi and other places for experimentation (Tan et al., 2012). The types of sample plots in the production area are diverse, including low mountains and hills, river valley plains and subtropical humid zones. The soil is mainly red soil and yellow soil, with an average annual temperature of 15℃ to 18℃ and an annual precipitation of 1200 to 1800 mm. In Changshan, Zhejiang Province, standardized cultivation and integrated pest and disease control are widely adopted. In newly introduced areas, adjustments are often made in combination with local traditional fruit tree management experience (Yu et al., 2012).

7.2 Research methodology: sampling, data collection, and comparative analysis

This case study established sample plots in places such as Changshan, Zhejiang Province and Chenzhou, Hunan Province. The zonal sampling method was adopted to select plants under different management models and systematically collect fruit samples (Tan et al., 2012; Yu et al., 2012; Huang et al., 2025). The sampling time covered stages such as fruit swelling, ripening and storage. The measured indicators included fruit size, weight, peel thickness, soluble solids, acidity, vitamin C content, as well as active components and volatile substances such as naringin and hesperidin (Gao et al., 2022; Miao et al., 2024; Wang et al., 2024; Ke et al., 2025). Meanwhile, by combining environmental factors such as meteorology, soil and management measures, and using multivariate statistics and principal component analysis methods, the fruit traits and yield performance of different ecological areas were compared (Tan et al., 2012; Yu et al., 2012).

7.3 Results: yield performance, fruit trait variations, and market responses

In the main production area of Changshan, Zhejiang Province, Changshan Huyou shows high and stable yield, with an annual yield of 50 to 80 kilograms per plant. The fruit size is uniform and the incidence of pests and diseases is low (Yu et al., 2012). In the newly introduced area of Chenzhou, Hunan Province, its adaptability is also relatively strong, the yield is stable, and the flavor is basically the same as that of the main production area. However, in some areas, due to differences in climate and soil, the maturity period may be slightly earlier or later (Tan et al., 2012). The average single fruit weight of the main production area is about 300 g. The fruit skin is relatively thick, the color is golden yellow, and the taste is a combination of sweet, sour and slightly bitter (Gao et al., 2022; Miao et al., 2024). The fruit size in the newly introduced area fluctuates, the content of soluble solids is slightly lower, and the acidity varies greatly, but the overall quality is still good (Tan et al., 2012). The fruit of the natural tetraploid Changshan Huyou is larger, the peel is thicker, and the content of some active components such as amino acids and flavonoids is higher, making it more suitable for medicinal use and deep processing (Huang et al., 2025). Changshan Huyou from Changshan, Zhejiang Province, has a strong market demand and stable prices due to its excellent quality and the fact that it is both edible and medicinal. Some products have even obtained geographical indication protection (Gao et al., 2022). Changshan Huyou in the newly introduced area is gradually being accepted by the market, but there is still room for improvement in brand building and quality standardization (Tan et al., 2012).

8 Challenges and Research Gaps

8.1 Lack of standardized trait evaluation across ecological zones

At present, there is no unified standard for the evaluation methods of the fruit traits of Changshan Huyou in different ecological regions. This makes it difficult to directly compare the research results from various regions, affecting the accuracy of variety breeding and regional adaptability analysis (Yu et al., 2012; Gao et al., 2022). There are significant differences in trait determination indicators, sampling time and evaluation systems among various regions (Tan et al., 2012). Meanwhile, there is a lack of a comprehensive evaluation system covering multiple aspects such as fruit quality, nutritional components and stress resistance (Gao et al., 2022). Some emerging technologies, such as molecular markers and metabolomics, have not been widely applied in regional trait evaluation (Miao et al., 2024; Huang et al., 2025). Due to the lack of unified standards, the research on variety promotion and regional adaptability lacks scientific basis, which hinders the precise breeding and promotion of superior varieties (Gao et al., 2022).

8.2 Trade-offs between yield quantity and fruit quality in different regions

In different ecological zones, high yield and high quality are often difficult to achieve simultaneously, and there is often a trade-off between yield and quality (Tan et al., 2012; Gao et al., 2022). Although some regions have high yields, the contents of soluble solids and flavor substances in the fruits have decreased, and the quality is not ideal (Huang et al., 2025; Ke et al., 2025). However, in areas with better fruit quality, the output is limited and the economic benefits are difficult to increase significantly (Chen et al., 2011; Tan et al., 2012). The influence mechanisms of factors such as climate, soil and management measures on yield and quality are very complex and have not been fully understood at present (Yu et al., 2012). This contradiction between yield and quality restricts the regionalized layout of varieties and the promotion of efficient cultivation models, and also affects the sustainable development of the industry (Gao et al., 2022; Ke et al., 2025).

8.3 Need for region-specific breeding and management recommendations

At present, the breeding and management methods of Changshan Huyou are mostly general, lacking customized plans for different ecological zones, which prevents the potential of the varieties from being fully exploited (Chen et al., 2011; Gao et al., 2022). Some regionally specific pests and diseases and adverse stress have not been fully incorporated into the breeding targets (Huang et al., 2025). Research on regionalized cultivation techniques, nutrient regulation and post-harvest treatment methods is also insufficient, and the promotion is not systematic enough (Tan et al., 2012). The regional trials and adaptability evaluation system of new varieties are still not perfect, and it is difficult for superior varieties to be precisely implemented (Gao et al., 2022). This insufficient regional adaptability has led to poor performance of some varieties when promoted in other places, thereby affecting industrial upgrading and farmers’ income (Chen et al., 2011; Huang et al., 2025).

9 Future Perspectives

9.1 Integrating ecological zoning with precision orchard management

Ecological zoning provides a scientific basis for regional adaptability in fruit tree production, while precision agriculture technology has brought orchard management into a data-driven and intelligent stage. In the future, the combination of these two aspects will promote the development of Changshan Huyou towards high efficiency and sustainability. By using remote sensing, the Internet of Things and intelligent sensors, key factors such as soil, water and climate can be monitored in real time, thereby optimizing strategies for fertilization, irrigation and pest control (Anastasiou et al., 2023; Soussi et al., 2024). For instance, decision-making systems based on big data and machine learning can provide tailor-made management plans for orchards in different ecological zones, improve the efficiency of resource utilization, and reduce the pressure on the environment at the same time (Finger et al., 2019; Júnior et al., 2024). Precise management can also make the size and quality of fruits more uniform, providing scientific support for processing and market circulation (Gao et al., 2022; Wang et al., 2024).

9.2 Breeding strategies for regional adaptability and quality optimization

The regional adaptability and fruit quality of Changshan Huyou are jointly influenced by genetics, environment and the interaction between the two. Future breeding strategies will place greater emphasis on genomic selection, molecular marker-assisted breeding and polyploid innovation (Szymański et al., 2020). The release of high-quality reference genomes provides a basis for identifying key genes that control adaptability, stress resistance and quality (Miao et al., 2024). Through systematic research on the differences in fruit traits in different ecological regions, germplasm resources with both strong adaptability and excellent quality can be screened out. Combined with molecular breeding technology, the breeding of new varieties can be accelerated (Guo et al., 2020). In addition, methods such as natural tetraploidization can not only make the fruits larger and more resilient, but also provide new directions for the development of characteristic products such as seedless citrus (Huang et al., 2025). Further research on the metabolic pathways of fruit nutrients and functional active substances is needed in the future to promote quality optimization and health value enhancement (Gao et al., 2022; Wang et al., 2024).

9.3 Outlook for industry competitiveness and regional development

With the formation of the Changshan Huyou industrial cluster, regional brand building and industrial chain extension have become the key points to enhance competitiveness. In the future, it is necessary to strengthen origin protection, promote standardized production, and enhance the added value and market influence of products through geographical indication certification (Yu et al., 2012). Meanwhile, efforts should be made to promote the integration of the primary, secondary and tertiary industries and develop emerging fields such as deep processing, functional foods and medicine, thereby enhancing the risk-resistance capacity and sustainability of the industries (Li et al., 2020a; Wang et al., 2024). In terms of regional economy, the development of Changshan Huyou industry can drive the agglomeration of related industries, increase farmers’ income and promote rural revitalization (Tan et al., 2012; Gao et al., 2022). In addition, it is necessary to attach importance to ecological environment protection and green development, promote the transformation of industries towards high efficiency, low carbon and circular, and achieve the coordinated improvement of economic, social and ecological benefits (Balafoutis et al., 2017; Finger et al., 2019; Petrović and Csambalik, 2025).

Acknowledgments

The authors appreciate the comments from the two anonymous peer reviewers on the manuscript of this study.

Conflict of Interest Disclosure

The authors affirm that this research was conducted without any commercial or financial relationships that could be construed as a potential conflict of interest.

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