1 Introduction

Ginkgo biloba L. is one of the oldest existing gymnosperms. It holds a unique position in evolution and has high research value. Its leaves and seeds contain many medicinal components, especially flavonoids and terpene lactones. These ingredients are widely used in the treatment of cardiovascular and cerebrovascular diseases and neurodegenerative diseases, and are globally best-selling herbal medicines and dietary supplements (Wang et al., 2023; Yao et al., 2023). In addition, the Ginkgo biloba has a unique leaf shape and its leaves turn golden in autumn, making it an important ornamental tree in gardens and very common in urban greening and landscape beautification (Han et al., 2023).

Ginkgo biloba has significant value both in medicine and ornamental purposes, but its genetic improvement and efficient breeding still face many challenges. Han et al. (2023) and Wang et al. (2023) demonstrated in their research that the life cycle of Ginkgo biloba is very long, with the juvenile stage possibly exceeding ten years, which slows down the progress of conventional breeding. Ginkgo biloba is dioecious and highly heterozygous. Its genetic background is very complex, making the stable inheritance and rapid aggregation of superior traits difficult (Yao et al., 2023). The genome of Ginkgo biloba is huge, and currently there is a lack of an efficient genetic transformation system. This has led to slow progress in molecular breeding and functional gene research, and limited the breeding efficiency of superior varieties.

This study summarizes the latest progress of Ginkgo biloba in medicinal and ornamental breeding, analyzes the main difficulties in genetic diversity, molecular marker development and phenotypic selection, discusses the application prospects of efficient breeding techniques, and proposes possible breakthrough directions for the future. The objective of this research is to provide a theoretical basis and technical support for the efficient utilization and industrial development of Ginkgo biloba resources.

2 Medicinal Value and Ornamental Importance of Ginkgo biloba

2.1 Key bioactive compounds (flavonoids, terpene lactones) and their pharmacological roles

Ginkgo biloba leaves and fruits contain many bioactive components, among which the most important ones are flavonoids and terpene lactones. These components have multiple pharmacological effects, including antioxidation, anti-inflammation, anti-cancer, antibacterial, antiviral, anti-platelet aggregation, and can also protect nerves and cardiovascular and cerebrovascular systems (Figure 1) (Mohanta et al., 2014; Šamec et al., 2022). Flavonoids have significant effects in regulating cardiac and cerebral blood flow, delaying neurodegenerative diseases, and improving cognitive function (Liu et al., 2022). Terpene lactones mainly play a role in regulating blood circulation, antioxidation and neuroprotection (Isah, 2015; Shahrajabian et al., 2021). In addition, Ginkgo biloba leaves and seeds contain various vitamins, minerals and polysaccharides, which also increase its medicinal value (Noor-E-Tabassum et al., 2022; Biernacka et al., 2023).

Figure 1 Bioactivity of G. biloba biflavonoids (Adopted from Šamec et al., 2022)

2.2 Ornamental traits: leaf morphology, autumn coloration, adaptability to urban environments

The leaves of Ginkgo biloba are fan-shaped and turn golden yellow in autumn, having a high ornamental value. Its leaf shapes are diverse. Some varieties can turn yellow in both spring and autumn and have a longer viewing period (Li et al., 2025). Ginkgo biloba can also tolerate pests and diseases, air pollution, drought and poor soil, and is very suitable for urban greening and street trees (Barker and Elston, 2022; Shareena and Kumar, 2022). Male plants are more popular in gardens because they have no unpleasant fruit smell. Ginkgo biloba has a long lifespan and often appears in historic cities and gardens. It is both a symbol of culture and a symbol of ecology (Isah, 2015; Šamec et al., 2022).

2.3 Dual-use breeding goals: balancing medicinal quality and ornamental value

Nowadays, the goal of Ginkgo biloba breeding is gradually shifting towards a balance between medicinal and ornamental uses. The focus of medicinal breeding is to increase the content of flavonoids and terpene lactones to enhance functions such as antioxidation and neuroprotection (Liu et al., 2022; Noor-E-Tabassum et al., 2022; Biernacka et al., 2023). Ornamental breeding focuses on leaf color, leaf shape, tree posture and environmental adaptability (Isah, 2015). The current difficulty lies in how to combine high medicinal components with excellent ornamental traits by using molecular breeding and genomic selection. In the future, more research is needed on the synthesis mechanism of medicinal components and the genetic basis of ornamental traits, so as to promote the comprehensive utilization of Ginkgo biloba in medicine and horticulture (Shareena and Kumar, 2022; Li et al., 2025).

3 Traditional Breeding Approaches

3.1 Selection and propagation of elite germplasm

The screening of superior germplasm has always been a core link in the traditional breeding of Ginkgo biloba. Wang et al. (2023) and Yao et al. (2023) established a core germplasm bank with strong representativeness through genetic diversity analysis of natural populations and cultivated varieties. These germplasm banks retain rich alleles and genetic diversity, providing important resources for the enhancement of medicinal components and the improvement of ornamental traits. Molecular markers have also been used to assist in germplasm identification and management, improving the recognition efficiency of superior genotypes and reducing resource duplication and homonymous foreign substances.

3.2 Hybridization attempts and limitations

Some progress has been made in hybrid breeding in the improvement of ornamental traits. Through artificial pollination and planned parent selection, a variety of new and superior varieties with stable leaf color and morphology have been obtained, some of which have performed outstandingly in garden and bonsai applications (Ming, 2001). However, the generation cycle of Ginkgo biloba is very long, the hybridization fruiting rate is low, and the trait segregation of offspring is large, resulting in an overly long hybridization breeding cycle, low efficiency, and difficulty in rapidly aggregating the target trait (Han et al., 2023).

3.3 Vegetative propagation and clonal selection for trait stabilization

Ginkgo biloba is propagated asexually through methods such as grafting and cuttings to avoid the separation of traits caused by sexual reproduction. Ming’s (2001) early research found that these techniques could achieve rapid propagation of superior individual plants and maintain the stability of traits. Clonal selection and breeding can ensure the consistency of medicinal components and ornamental traits, and also facilitate large-scale promotion. Han et al. (2023) found that in recent years, molecular biology and cell engineering techniques have provided new tools for the study of Ginkgo biloba gene functions and the optimization of asexual reproduction systems, which are expected to further enhance the stability of superior traits and the efficiency of genetic improvement.

4 Molecular and Genomic Breeding Strategies

4.1 Development of molecular markers (SSR, SNP, AFLP) for trait mapping

Researchers have developed a large number of markers such as SSR, SNP and InDel by using transcriptome and genomic data. Based on the studies of EST-SSR and InDel, it was found that the Ginkgo biloba germplasm resources have a high genetic diversity, and a core germplasm bank was successfully established. These achievements provide basic tools for trait mapping and genetic improvement (Wang et al., 2023; Yao et al., 2023). Wu et al. ’s research in 2019 demonstrated that the development of SNP markers and the application of high-resolution melting curve technology have also enhanced the efficiency of genetic diversity analysis and functional gene mining in Ginkgo biloba populations. These molecular markers have been widely used in fingerprint mapping construction, germplasm identification, genetic diversity evaluation and core germplasm screening.

4.2 Genomic insights: transcriptomics, genome sequencing, and gene mining

Han et al. (2023) demonstrated that the draft whole genome and high-quality genome of Ginkgo biloba have been released, providing a scientific basis for the mining of functional genes and the study of complex traits. Transcriptome sequencing revealed the key genes and their regulatory networks related to the synthesis of important secondary metabolites such as flavonoids and lignin (Wu et al., 2018). Some transcription factors have been systematically identified and expression analyzed, providing target genes for increasing the content of medicinal components and improving stress resistance (Zhou et al., 2020). Genome-wide gene family studies have expanded the molecular understanding of Ginkgo biloba growth and development and stress response (Guo et al., 2023; Li et al., 2024).

4.3 Marker-assisted selection (MAS) and genomic selection prospects

By combining molecular markers and phenotypic data, target traits (such as medicinal component content, leaf color, stress resistance, etc.) can be screened more efficiently, and selection can be made at an early stage. Ginkgo biloba has a long life cycle and a complex genetic background. However, both MAS and genomic selection (GS) have shown great potential in core germplasm construction, aggregation of superior genotypes, and conservation of genetic diversity (Wang et al., 2023; Yao et al., 2023). With the increase in the number and coverage of molecular markers and the in-depth study of functional genes, in the future, MAS and GS are expected to accelerate the breeding process of new Ginkgo biloba varieties and achieve targeted improvement of medicinal and ornamental traits (Wu et al., 2019).

5 Biotechnological Approaches

5.1 Tissue culture, somatic embryogenesis, and micropropagation

Tissue culture and somatic embryogenesis provide an important foundation for the rapid reproduction and preservation of superior genotypes of Ginkgo biloba. The cell culture system of Ginkgo biloba has been used to screen and optimize cell lines with high-yield medicinal components. The yield of bioactive substances can be increased by adjusting the culture conditions and inducing differentiation and other methods. Micropropagation technology can also solve problems such as long growth cycle and low propagation efficiency of Ginkgo biloba, providing technical support for large-scale production of high-quality plants and genetic improvement (Sabater-Jara et al., 2013).

5.2 CRISPR/Cas and gene-editing possibilities for trait improvement

Although the genome of Ginkgo biloba is large and its genetic transformation system is not yet perfect, progress has been made in recent years. Researchers have established a protoplast isolation and transient expression system, providing a scientific basis for gene function research and gene editing. Efficient protoplast isolation and PEG-mediated transient transformation system make it possible to study subcellular localization, overexpression and protein-protein interaction of genes, providing a technical platform for the application of gene editing tools such as CRISPR/Cas in the improvement of Ginkgo biloba traits in the future (Han et al., 2023).

5.3 Metabolic engineering for enhanced medicinal compound production

The flavonoids and terpene lactones in Ginkgo biloba leaves are the main medicinal components. Through the combined analysis of transcriptome and metabolome, researchers have identified the key genes and transcription factors regulating flavonoid biosynthesis, providing targets for metabolic engineering (Wu et al., 2018; Guo et al., 2020). Meanwhile, optimizing the cell culture system, applying exogenous inducers, and regulating related genes can significantly increase the accumulation of medicinal components in Ginkgo biloba cells (Sabater-Jara et al., 2013). These methods provide new ideas for the sustainable development and high-value utilization of Ginkgo biloba medicinal resources.

6 Case Study: Breeding Program for Improved Ginkgo biloba

6.1 Selection of high-flavonoid lines and ornamental cultivars

Ginkgo biloba is a tree species that has both medicinal and ornamental value. Its breeding goals mainly focus on increasing the content of flavonoid active components in leaves and improving ornamental traits. Research has found that there are significant differences in the content of flavonoids (such as isorhamnetin) in the leaves of Ginkgo biloba with different crown types. Medium-sized crown-shaped individuals with a larger crown width not only have strong growth vigor and a higher leaf volume, but also have a higher flavonoid content. Therefore, through phenotypic screening of the crown type, the content of active components of Ginkgo biloba can be effectively increased, providing a basis for the breeding of medicinal Ginkgo biloba (Wu et al., 2020). In terms of ornamental breeding, four new ornamental leaf varieties have been obtained through sexual hybridization, and approximately 11 000 stable seedlings have been cultivated, enriching the variety resources of gardens and bonsai (Ming, 2001).

6.2 Methods used: marker analysis, hybridization, and phenotypic screening

Molecular markers have been used to analyze and manage germplasm resources, reveal genetic diversity, and help establish core germplasm banks. These achievements provide a basis for molecular-assisted selection and genetic improvement (Wang et al., 2023; Yao et al., 2023). Hybrid breeding has obtained new materials with excellent traits through the selection of superior parents, artificial pollination and seedling screening (Ming, 2001). Phenotypic screening mainly focused on morphological indicators such as crown shape, leaf area, and leaf density, and combined with the detection of active components, achieved simultaneous improvement of medicinal and ornamental traits (Wu et al., 2020). Han et al. (2023) demonstrated that the establishment of a protoplast isolation and transient transformation system provides a new platform for the study of Ginkgo biloba gene functions and molecular breeding.

6.3 Outcomes: improved cultivars and implications for large-scale cultivation

Through these strategies, researchers have obtained a batch of new Ginkgo biloba varieties with high flavonoid content and excellent ornamental traits. These varieties have enhanced the quality and output of medicinal raw materials and also expanded the options for landscaping and landscape beautification. The establishment of core germplasm banks and molecular marker-assisted selection have effectively enhanced breeding efficiency and germplasm management level, laying the foundation for large-scale promotion and industrialization (Wang et al., 2023; Yao et al., 2023). The new varieties obtained through hybrid breeding also show broad application prospects in the garden and bonsai markets (Ming, 2001). With the continuous progress of molecular biology and gene function research, Ginkgo biloba breeding will become more precise and efficient, promoting its development in the fields of medicine and ornamental (Han et al., 2023; Wang et al., 2023).

7 Challenges and Limitations

7.1 Long juvenile phase and dioecy complicating breeding cycles

The juvenile period of Ginkgo biloba is very long, usually taking more than ten years to flower and bear fruit. This greatly prolongs the breeding cycle and also reduces the efficiency of new variety selection and breeding. In addition, Ginkgo biloba is dioecious, and it is difficult to distinguish between female and male plants in the early stage. During artificial hybridization, strict control of pollination targets is also required, which makes the breeding process more complicated and prolongs the cycle (Ming, 2001; Han et al., 2023).

7.2 Limited genomic resources and functional validation tools

In recent years, some progress has been made in the sequencing of Ginkgo biloba genomes. However, the large and complex genomic structure, limited molecular markers, and the lack of an efficient genetic transformation system have restricted the research on functional genes and the development of molecular breeding. Most current studies rely on heterologous systems to verify gene functions, but the reliability and applicability of this approach are limited. The newly established protoplast transient expression system provides a new platform for functional validation, but still cannot replace the stable genetic transformation system (Han et al., 2023; Wang et al., 2023; Yao et al., 2023).

7.3 Trade-offs between medicinal quality and ornamental traits

The medicinal value of Ginkgo biloba mainly comes from the active components such as flavonoids and terpene lactones in its leaves, while its ornamental properties are mainly concentrated on the appearance features such as leaf color and leaf shape. Some ornamental traits (such as special leaf colors) may be negatively correlated with the accumulation of active ingredients, which makes it difficult to balance medicinal quality and ornamental traits simultaneously. In addition, there are significant differences in genetic basis between superior medicinal varieties and ornamental varieties, which also increases the difficulty of collaborative improvement (Wang et al., 2023; Li et al., 2025).

8 Future Perspectives

8.1 Application of multi-omics and big data in Ginkgo breeding

With the development of genomics, transcriptomics and metabolomics, people have gained new insights into the genetic basis and functional genes of Ginkgo biloba. High-throughput sequencing and genotyping techniques have been employed to analyze the genetic diversity and population structure of Ginkgo biloba germplasm resources and to establish core germplasm banks. These achievements have laid the foundation for molecular breeding of medicinal and ornamental traits (Wang et al., 2023). Wang et al. (2023) and Yao et al. (2023) can more accurately locate functional genes, screen out superior genotypes, and accelerate the breeding process of high-quality varieties through the integration and analysis of genetic information based on big data.

8.2 Integrating molecular tools with conventional selection for efficiency

Traditional hybridization, selection and grafting methods are crucial in improving the ornamental traits and enhancing the medicinal components of Ginkgo biloba, but these methods are limited by the long generation cycle and complex genetic background of Ginkgo biloba. In recent years, molecular marker-assisted selection (MAS) and gene function analysis platforms have provided new tools for the early screening of target traits and gene function verification (Han et al., 2023; Wang et al., 2023; Yao et al., 2023). The combination of molecular tools and conventional breeding methods can enhance efficiency, shorten the cycle and improve the accuracy of new variety breeding.

8.3 Breeding for climate-resilient, dual-purpose cultivars

In the face of climate change and diverse application demands, the future goal is to cultivate new varieties of Ginkgo biloba that possess both medicinal and ornamental value as well as climate adaptability. Genetic diversity analysis and the establishment of core germplasm banks can provide abundant genetic resources for the synergistic improvement of stress resistance, medicinal components and ornamental traits (Wang et al., 2023; Yao et al., 2023). By using molecular markers and multi-omics data to screen and aggregate genotypes with stress resistance, high quality and high ornamental value, it is expected to breed new Ginkgo biloba varieties that can adapt to different ecological environments and meet diverse market demands (Li et al., 2025).

Acknowledgments

The authors appreciate the modification suggestions from two anonymous peer reviewers on the manuscript of this study. The authors also thank the group members for their assistance during the research process.

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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