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Review

Unlocking the Potential of Sophora moorcroftiana (Fabaceae): The Overlooked Xizang Endemic

by
Duozhuoga Mei
1,
Sinong Yu
2,
Shuangyuan Yu
1,
Fuliang Cao
1,
Guibin Wang
1 and
Tingting Dai
1,*
1
Co-Innovation Center for Sustainable Forestry in Southern China, Nanjing Foresty University, Nanjing 210037, China
2
Modern Forestry Innovation Center of Yancheng State-Owned Forest Farm, Yancheng 048000, China
*
Author to whom correspondence should be addressed.
Forests 2025, 16(3), 410; https://doi.org/10.3390/f16030410
Submission received: 27 January 2025 / Revised: 19 February 2025 / Accepted: 20 February 2025 / Published: 24 February 2025
(This article belongs to the Section Forest Ecology and Management)
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Abstract

:
Sophora moorcroftiana (Benth.) Baker is a drought- and sand-resistant endemic shrub species in the family Fabaceae, native to the Tibetan Plateau along the Yarlung Tsangpo River (elevation: 2800–4400 m). This study offers a comprehensive review of the latest research on S. moorcroftiana, with a focus on its ecological functions, medicinal potential, pest and disease management, and germplasm conservation. By synthesizing existing studies, the review sheds light on the mechanisms that enable this species to thrive in extreme environments, highlights its unique secondary metabolites, and explores its critical role in biodiversity maintenance. Additionally, the article examines the current conservation status of S. moorcroftiana, identifies the key threats to its survival, and suggests future research directions and strategies for sustainable utilization. The goal of this review is to fill existing knowledge gaps by providing a theoretical foundation and practical guidance for future scientific research, applied uses, and conservation initiatives related to S. moorcroftiana.

1. Introduction

Sophora moorcroftiana (Benth.) Baker, a member of the genus Sophora within the family Fabaceae, is a perennial dwarf shrub typically ranging in height from 1 to 3 m (Figure 1). Its dense branches often culminate in sharp, rigid spines that bear a striking resemblance to wolf teeth, serving both a defensive and structural role. As a result, the species is commonly referred to as “Wolf Tooth Thorn” [1]. The inflorescence of S. moorcroftiana is racemose, appearing at the tips of twigs or within leaf axils. The calyx is bell-shaped and exhibits a blue hue, while the corolla displays a vibrant blue-purple color, which has earned the plant the popular name “Lavender of the Plateau” [2,3,4]. The pods present a beaded appearance and dehisce along their inner suture, releasing smooth, light yellow-brown seeds that are ovoid to spherical in shape. These seeds measure approximately 4.5 mm in length and 3.5 mm in diameter [5]. The flowering period spans from May to July, with fruiting occurring between July and October [1,6,7].
This species is primarily distributed in the middle and lower reaches of the Yarlung Tsangpo River Basin, encompassing regions such as Nyingchi, Lhasa, Shannan, Shigatse, and others. A specialized survey by Ning Xiaobin et al. [8] on S. moorcroftiana resources in Xizang revealed that the species occupies an area of 206,560 hectares across 32 counties (districts) and 160 townships within five prefectures: Lhasa, Shannan, Shigatse, Nyingchi, and Ngari (Figure 2). Within these regions, Samzhubze District in Shigatse holds the largest distribution area at 34,623.14 hectares, followed by Lhatse County with 19,943.11 hectares. In contrast, Bayi District in Nyingchi has the smallest distribution, covering only 3.07 hectares. Additionally, S. moorcroftiana is also found in neighboring countries such as India, Bhutan, and Nepal.
S. moorcroftiana is particularly renowned for its secondary metabolites, especially its notably high total alkaloid content compared to other species. The market value of matrine, one of its primary alkaloids, can reach up to 2.6 million RMB per ton [9], earning the plant the moniker “Gold of the Xizang Plateau”. Extracts from this species are extensively utilized in the pharmaceutical industry due to their antibacterial [10], antiviral, and anticancer properties [11]. Beyond its medicinal applications, the leaves of S. moorcroftiana are an important resource for fodder and firewood in Xizang [12].
However, the intensification of human activities and habitat destruction has severely threatened the natural populations of S. moorcroftiana. In regions where grazing, land reclamation, and infrastructure development are prevalent, the species’ population is gradually declining and may even face the risk of local extinction. Therefore, the conservation and collection of S. moorcroftiana germplasm resources are of paramount importance for ecological restoration and the preservation of genetic diversity. Such efforts not only contribute to mitigating ecological issues such as desertification and land degradation but also provide valuable material for future research.
The primary objectives of this review are to (1) synthesize the latest research on the ecological functions, medicinal potential, and conservation status of S. moorcroftiana; (2) identify the key mechanisms enabling its adaptation to extreme environments; and (3) provide practical recommendations for sustainable utilization and conservation strategies. By addressing these goals, this review aims to fill existing knowledge gaps and support future research and conservation efforts.

2. Ecological and Environmental Value

S. moorcroftiana plays a critical role in the ecological balance of the Qinghai Xizang Plateau, serving as an indispensable component of the region’s ecosystem. Through its unique adaptability and specialized physiological characteristics [13], S. moorcroftiana not only thrives in the harsh, high-altitude, arid, and barren environments of the plateau but also contributes to regional ecosystem stability through several key mechanisms, including nitrogen fixation, water conservation, and the enhancement of soil fertility and structure [14]. As a natural windbreak and sand binding plant, it has proven to be highly effective in combating land desertification, thereby safeguarding agricultural and pastoral activities, as well as human habitation. Additionally, S. moorcroftiana provides vital habitats and shelters for various plants and animals, thus playing a significant role in enhancing local biodiversity [14]. Its remarkable drought tolerance and salt-alkali resistance further establish it as an irreplaceable pioneer species in ecological restoration projects, particularly those focused on soil erosion control and the rehabilitation of degraded grasslands [15]. These ecological functions highlight S. moorcroftiana’s importance not only as a highly adaptable native plant but also as a key factor in maintaining the health and sustainability of the Qinghai Xizang Plateau ecosystem. It is recognized as one of the region’s most resilient shrub species, known for its resistance to solar radiation and sand burial, and is considered a pioneer species in studies of adaptation to changing climatic conditions [16].

2.1. Windbreak and Sand Stabilization in Combating Desertification

In recent years, the expansion of sandy areas and the degradation of vegetation on the Xizang Plateau have become increasingly prominent. As a pioneer species uniquely adapted to sandy environments, S. moorcroftiana demonstrates exceptional ecological adaptability. Its dual reproductive mechanisms, seed propagation, and root sprouting, enable it to establish initial vegetation on shifting sand dunes and maintain growth in stabilized sandy areas [17]. The introduction of S. moorcroftiana has significantly improved soil fertility and water retention. After its introduction, the soil organic matter content increased from 0.13 g/kg to 9.49 g/kg, a remarkable 72-fold increase, and the total nitrogen content rose from 0.02 g/kg to 0.66 g/kg, a 32-fold increase. In the 10 cm soil layer, the saturated water content and capillary water content increased by 52.65% and 88.75%, respectively, and the soil evaporation rate decreased by 37.86%–62.24%. These significant improvements in soil quality not only stabilize sand dunes but also enhance ecosystem stability by increasing soil organic matter and improving hydrological functions [18].
S. moorcroftiana plantations have significant ecological advantages in windbreak and sand stabilization. The canopy, litter accumulation, and biological crusts of the plants work together to reduce soil moisture loss by limiting surface evaporation. The soil hydrological functions of S. moorcroftiana plantations are 1.71–2.62 times stronger than those of active sand dunes, primarily due to improvements in soil organic matter content and water holding capacity [19]. Although sand burial can reduce the density of the seed bank and the number of seedlings of S. moorcroftiana, studies have shown that sand burial can stimulate the sprouting and growth of branches, highlighting the species’ adaptability and positive response to sand burial environments [16,20].
With the advent of global climate change, the Qinghai Tibet Plateau region has witnessed a significant increase in temperature and alterations in precipitation patterns [21], which have far-reaching implications for S. moorcroftiana. In the short term, a rise in temperature might potentially expand the habitat of S. moorcroftiana and facilitate its growth and dispersal. However, when the temperature continuously increases beyond its adaptability threshold, the seed germination rate and seedling survival rate are likely to decline [22]. This will make it more difficult to establish vegetation on sand dunes. Regarding its suitable growth environment, it thrives at an altitude ranging from 3400 to 4250 m, with an isothermality of 43–48 and the mean temperature of the coldest month within the range of −19 °C to −9 °C [22]. Under future climate scenarios, the core distribution area of S. moorcroftiana will shift westward along the Yarlung Tsangpo River Basin. High temperatures will limit its optimal growth conditions in cold environments, leading to a notable contraction of its habitat range.
Moreover, although S. moorcroftiana is highly drought resistant and can stabilize the soil in arid and sandy environments with its deep root system, changes in precipitation patterns and prolonged droughts will exacerbate soil moisture limitations, further weakening its ecological restoration and sand-fixing functions. Extreme rainstorms may wash away its roots, undermining the soil holding capacity and intensifying soil erosion [23,24]. Meanwhile, long-term droughts will affect the activity of soil microorganisms and alter the structure of the rhizosphere microbial community, which is detrimental to the improvement of soil fertility [25].
Therefore, against the backdrop of future climate change, the conservation and artificial cultivation of S. moorcroftiana are of great significance. These efforts can provide a scientific basis and theoretical guidance for windbreak, sand fixation, and desertification control practices.

2.2. Soil and Water Conservation and Fertility Enhancement

Boasting an extensive root system and abundant foliage, S. moorcroftiana plays a critical role in mitigating soil erosion and improving soil structure. The productivity of S. moorcroftiana is significantly influenced by its age and environmental parameters such as soil depth, moisture, and pH. Studies have shown that as the plant ages, its efficiency in utilizing and accumulating nutrients like nitrogen and phosphorus changes, directly affecting its growth and recovery capacity. For instance, mature plants exhibit lower nutrient use efficiency in arid, nutrient-poor environments, which limits their productivity [26,27]. Additionally, environmental factors such as soil total phosphorus and pH significantly influence plant growth at different restoration stages, highlighting the role of temporal dynamics in regulating productivity [22]. Therefore, understanding the interaction between these environmental factors and plant age is crucial for accurately predicting the recovery potential of S. moorcroftiana. Its foliage intercepts rainfall, reduces evaporation, and enhances soil organic matter content, thereby indirectly boosting soil fertility. By stabilizing soil and increasing erosion resistance, S. moorcroftiana is recognized as a keystone species in combating soil degradation and land desertification [26].
This species exhibits remarkable ecological adaptability to the arid regions of the Xizang Plateau where its growth is strongly influenced by soil nutrient availability (e.g., potassium, phosphorus, and total nitrogen) and topographic factors (slope aspect and altitude) [27]. Research has revealed that S. moorcroftiana exhibits distinct growth patterns in different soil types. For example, its biomass positively correlates with potassium and phosphorus levels in riverbank sandy soils, while total nitrogen content becomes a limiting factor in hillside sandy soils [26,27]. Furthermore, as soil depth increases, the contents of soil organic carbon (SOC), total nitrogen (TN), total phosphorus (TP), and total potassium (TK) show an upward trend, whereas soil pH decreases, indicating the significant influence of soil layers on nutrient accumulation [22]. These findings demonstrate that the availability of soil nutrients directly affects the growth and productivity of S. moorcroftiana, thereby influencing its role in ecological restoration.
For example, in riverbank sandy soils, its biomass accumulation positively correlated with potassium and phosphorus levels, whereas on hillside sandy soils, total nitrogen content emerges as the limiting factor. Furthermore, S. moorcroftiana facilitates the accumulation of soil organic matter and enhances carbon, nitrogen, and phosphorus storage, making it indispensable for ecosystem restoration. While it stabilizes sandy soils and slows weathering, nutrient deficiencies, particularly in nitrogen, remain critical constraints on its population health, underscoring the need for targeted soil fertility management [28].
The rhizosphere microorganisms of S. moorcroftiana are integral to soil fertility enhancement. Notably, plant growth-promoting rhizobacteria (PGPR) enhance nutrient utilization efficiency, improve stress tolerance, and optimize soil conditions [29]. Moreover, the rhizosphere microbial community plays a pivotal role in improving soil fertility. Studies have shown that specific plant growth-promoting rhizobacteria (PGPR) significantly enhance the availability of nitrogen and potassium, thereby optimizing plant growth conditions [21]. For instance, the PGPR strain SH505 not only increases nitrogen and potassium availability but also boosts soil enzyme activities such as cellulase, sucrase, and catalase, thus improving soil conditions [22]. Additionally, the composition and diversity of soil microbial communities change with increasing planting age, which in turn affects the multifunctionality of soil ecosystems [22]. Therefore, understanding the interactions between rhizosphere microorganisms and plants is essential for optimizing soil management and enhancing plant productivity.
Among these, the PGPR strain SH505, isolated from S. moorcroftiana’s rhizosphere, demonstrates exceptional potential: it elevates available nitrogen and potassium levels, stimulates cellulase, sucrase, and catalase activities, and modulates soil pH and electrical conductivity. With its robust tolerance to salinity, drought, and extreme pH, SH505 represents a promising candidate for microbial fertilizer development [30].
In China’s intensive agricultural systems, despite high fertilizer inputs, nitrogen use efficiency remains alarmingly low at 19%. This inefficiency leads to accumulation, nitrogen waste, soil acidification, and increased susceptibility to soil-borne pathogens, ultimately compromising crop yields and agricultural sustainability [31]. In this context, nitrogen-fixing bacteria offer a sustainable solution by converting nitrate into bioavailable forms, thereby improving use efficiency and soil fertility [32].
As a leguminous plant, S. moorcroftiana forms symbiotic root nodules with nitrogen-fixing bacteria. Recent studies in the Yarlung Tsangpo River Basin have identified 16 PGPR strains in its rhizosphere, two of which exhibit high nitrogen-fixing capacity [33]. Interestingly, S. moorcroftiana shows remarkable resistance to pathogenic fungi, likely due to the protective functions of its rhizosphere microbiome [34]. Its growth is significantly influenced by age and environmental parameters, and understanding these relationships is essential for predicting recovery and developing effective soil management strategies. For example, the availability of soil nutrients such as nitrogen, phosphorus, and potassium, as well as the composition of microbial communities, directly impacts plant productivity, while environmental factors like soil total phosphorus and pH further regulate plant growth at different restoration stages [19,20]. Future research should focus on the interactions between microbial communities and plant growth, as well as the changes in soil ecosystem functions under different planting ages, to further improve the success of ecological restoration.

2.3. Genetic Regulation of Drought Stress Responses in Sophora moorcroftiana

S. moorcroftiana activates a coordinated genetic network to combat drought stress (Table 1). Central to this response are dehydration-responsive element-binding proteins (DREBs), whose expression is significantly upregulated under drought conditions [35]. Comparative analyses with model plants reveal evolutionary divergence: For instance, Arabidopsis thaliana AtDREB1A operates via ABA-independent pathways [36], whereas S. moorcroftiana DREBs (SmDREB1–4) likely employ more complex regulatory mechanisms, a subject requiring further investigation. Among these, SmDREB1 is uniquely activated by drought, cold, and heat stress, with peak expression in leaves. Unlike the cold-specific AtDREB1/CBFs [37,38,39], SmDREB1 shares functional parallels with Medicago truncatula MtDREB1C, which confers drought/salt tolerance via ABA-independent pathways [40]. Mechanistically, SmDREB1 induces AMY3 (α-amylase), triggering starch-to-sugar conversion to enhance osmotic adjustment and carbon partitioning during drought [41].
In contrast, SmDREB2 is predominantly expressed in roots and flowers under salt, cold, and heat stress, mirroring the phosphorylation-dependent activation of Arabidopsis DREB2A [42]. Similarly, SmDREB3 and SmDREB4 are upregulated by drought, heat and salt, cold, and heat stress, respectively, suggesting specialized roles in stress adaptation.
These DREB proteins synergize with MYB transcription factors, e.g., (MYB2 and MYB43) to amplify drought-responsive gene expression [43]. Additionally, aquaporins (AQPs) and dehydrins (DHNs) maintain cellular water balance, while antioxidant enzymes (e.g., SOD, POD, and CAT) scavenge reactive oxygen species (ROS). Abscisic acid (ABA) signaling further modulates this response, with ABA-responsive genes contributing to enhanced drought tolerance. Functional validation via SmDREB1 overexpression in Arabidopsis confirmed its role in drought adaptation: transgenic lines exhibited elevated AtDHN, AtLEA, AtPIP2;2, AtPIP2;3, and AtRD29 expression, accompanied by higher leaf water potential, antioxidant activity, proline accumulation, and sustained photochemical efficiency (Fv/Fm) under drought [44].
Genomic studies [45] highlight 11 sucrose metabolism genes (e.g., AMY3 and CWINV1) as critical for drought adaptation. Root-specific sucrose accumulation, driven by MY3-mediated starch hydrolysis and CWINV1-regulated sucrose cleavage, enhances osmotic adjustment and carbon allocation [41,46]. Supporting this, seed osmopriming combined with biochar application boosts water use efficiency (95%), carbon assimilation, and antioxidant capacity (34% reduction in MDA), significantly alleviating drought impacts [47].
In conclusion, S. moorcroftiana drought resilience stems from the synergistic action of stress-responsive genes, optimization, and ROS scavenging systems. These insights provide a molecular framework for engineering drought-tolerant crops.
Table 1. List of drought resistance-regulating genes in S. moorcroftiana.
Table 1. List of drought resistance-regulating genes in S. moorcroftiana.
GenesDescriptionGene_IDFunctionRefs.
SmDREB1Drought-responsive element-binding protein 1KM527092Regulates downstream drought resistance genes[22,43,44]
SmDREB2Drought-responsive element-binding protein 2KM527093Protects cells and helps retention[22]
SmDREB3Drought-responsive element-binding protein 3KM527094Protects cells[22]
SmDREB4Drought-responsive element-binding protein 4KM527095Facilitates water transport[22]
AMY3Alpha-amylaseSsh0029976Hydrolyzes starch to release sugars[45]
CWINV1Beta-fructofuranosidaseSsh0033248Hydrolyzes sucrose[45]

3. Medicinal Value

3.1. Active Compounds and Clinical Applications

The seeds of S. moorcroftiana are rich source of bioactive alkaloids, including matrine, oxymatrine, sophocarpine, and oxysophocarpine [48]. These compounds exhibit a wide range of pharmacological activities, making them valuable traditional Xizang and Chinese medicine for their anti-inflammatory, anticancer, antiviral, antioxidant, and immunomodulatory properties.
Among these alkaloids, oxymatrine is particularly notable for its potent anti-inflammatory effects. Oxymatrine can alleviate symptoms of ulcerative colitis (UC) by modulating immune cell functions and inhibiting the NF-κB pathway, This suppresses of the NF-κB pathway lead to reduced expression of pro-inflammatory cytokines (e.g., TNF-α and IL-6) and alleviation of intestinal inflammation [49,50]. Beyond its anti-inflammatory properties, oxymatrine also significant anti-tumor activity. It induces apoptosis in cancer cells through the mitochondrial pathway, upregulating pro-apoptotic proteins (e.g., Bax) while downregulating anti-apoptotic proteins (e.g., Bcl-2). These mechanisms contribute to its efficacy in suppressing the growth of various cancer cells, including cervical cancer [51].
Matrine and sophocarpine further enhance the therapeutic potential of S. moorcroftiana. Matrine exerts anticancer effects by inhibiting the PI3K/AKT/mTOR signaling pathway, which regulates cell proliferation and survival, while also providing protection against oxidative stress and inflammation in organs such as the liver and intestines [52,53]. Sophocarpine complements these effects by inducing cell cycle arrest and activation caspase-3 and caspase-9 [54]. The synergistic action of these alkaloids is evident in total alkaloids extracted (TA-SM), which exhibit strong anti-angiogenic activity by signaling VEGF and inhibiting the proliferation and migration of vascular endothelial cells [55]. Moreover, water soluble alkaloids, such as E2 (a combination of matrine and sophocarpine), have shown promise in treating UC due to their robust anti-inflammatory effects [50,56]. In addition to their anticancer and anti-inflammatory properties, S. moorcroftiana extracts demonstrate significant antibacterial activity. They are effective against methicillin resistant Staphylococcus aureus (MRSA), potentially addressing drug resistance through mechanisms such as cell wall disruption and the inhibition of bacterial efflux pumps [11]. This multifaceted pharmacological profile underscores the therapeutic versatility of S. moorcroftiana alkaloids.
The complexity of these alkaloids’ mechanisms of action has been further elucidated through network pharmacology. For example, the drug CIPHER algorithm predicted that matrine acts on a network of targets involved in neurotoxicity, neuropharmacological modulation, macropinocytosis induction, and ATP metabolism regulation [57]. Experimental studies confirmed these predictions, showing that matrine induces macropinosome formation and reduces cellular ATP levels, highlighting the value of network pharmacology in uncovering the systemic effects of natural products.
Advances in extraction technologies have also played a crucial role in optimizing the isolation and quantification of these bioactive compounds. Traditional methods, which often suffer from poor selectivity and high solvent consumption, have been improved through the development of double-templated molecularly imprinted polymers (D-MIP) combined with HPLC-MS/MS [58,59,60,61]. This approach, using matrine and oxymatrine as template molecules, achieves high selectivity and sensitivity, with recovery rates ranging from 73.25% to 98.42% and detection limits as low as 9.23–15.42 μg/kg. This method not only reduces matrix interference but also offers a fast and environmentally friendly solution for extracting and analyzing matrine-type alkaloids in S. moorcroftiana and related species [62]. These advancements highlight the potential for the scalable and efficient extraction of bioactive compounds from natural sources.
Environmental factors such as altitude, latitude, and longitude significantly influence the alkaloid content of S. moorcroftiana. Studies have shown that oxymatrine levels are positively correlated with altitude, while matrine content decreases with higher latitude and longitude [48]. These findings provide valuable insights for selecting high-yield populations and optimizing cultivation strategies to maximize the production of these pharmacologically important metabolites.

3.2. Safety of Consumption

While the alkaloid components of S. moorcroftiana have shown remarkable pharmacological effects, their safety remains a key focus of ongoing research. Compounds such as oxymatrine and matrine exhibit certain toxic effects, especially at high doses, potentially affecting the liver, nervous system, and heart. For instance, studies in mouse models have indicated that high doses of oxymatrine and matrine may lead to liver dysfunction, neurotoxicity, and even developmental toxicity [53,63]. Oxymatrine may cause mild liver dysfunction and hematotoxicity, underscoring the need for stringent dosage control in clinical applications [63]. Similarly, sophocarpine at high doses may lead to cardiotoxic effects, although its overall toxicity is relatively low [64,65].
Toxicity assessments have shown that matrine and oxymatrine are generally less toxic to normal cells, with oxymatrine demonstrating favorable selectivity for treating certain diseases. However, the long-term safety and potential risks associated with high-dose usage require further investigation. For example, the prolonged use of TA-SM may affect other physiological systems, necessitating comprehensive safety assessments in clinical applications [5].
To ensure safe usage, regulatory guidelines for S. moorcroftiana-based preparations should be established, including maximum allowable doses and contraindications for specific populations (e.g., pregnant women and individuals with liver or heart conditions). Furthermore, future research should focus on developing standardized extraction methods and formulations to minimize toxicity while maximizing therapeutic efficacy.

4. Pests and Disease Management

4.1. Major Pests and Diseases

In recent years, seed pests in leguminous shrub forests have emerged as a critical constraint to the industrial development of S. moorcroftiana. The cryptic feeding behavior and high destructiveness of Etiella zinckenella (Treitschke) and the Bruchophagus onois (Mayr) (Figure 3) make them particularly challenging to manage, threatening both natural regeneration and artificial cultivation.

4.1.1. Pod Borer (Etiella zinckenella)

The pod borer (Etiella zinckenella), a lepidopteran pest, primarily targets leguminous plants, feeding on pods during its larval stage. Adult moths, equipped with strong flight abilities, lay their eggs on pod surfaces. Upon hatching, the larvae penetrate the pod wall and consume the seeds, with their feeding activity concentrated mainly in the upper canopy. Larval feeding increases seed moisture content and contaminates the seeds with excrement and decayed tissue, leading to a substantial decline in seed quality. In cases of severe infestation, seeds may become completely non-viable, with germination rates dropping to zero [66]. Furthermore, the larvae cause pod discoloration and softening, further compromising seed integrity. Recent studies by Zang Jiancheng et al. [19] on the mitochondrial genome of E. zinckenella populations in the Lhasa region revealed a close phylogenetic relationship to a novel species within the genus Meroptera, suggesting a potential evolutionary link within the order Lepidoptera. This finding lays a molecular foundation for future population monitoring and pest management efforts.

4.1.2. Seed Wasp (Bruchophagus onois)

The seed wasp (Bruchophagus onois), a hymenopteran species, lays its eggs within the seeds of S. moorcroftiana, where the larvae feed on the endosperm, causing substantial damage to the seed structure. The first-generation larvae typically consume the entire seed endosperm, while the second-generation larvae partially consume it and overwinter within the seed. Infested seeds exhibit significantly reduced germination rates, averaging only 9.4%. The cumulative damage to seeds can have lasting effects on the species’ regeneration potential [67]. In extreme infestations, seed infestation rates may reach as high as 70%, presenting serious threats to both the natural regeneration and artificial cultivation of S. moorcroftiana.
The ecological impacts of these pests extend beyond direct seed damage. Field studies indicate that pest infestation rates are influenced by altitude and microclimate, with higher infestation rates observed in low-altitude shrublands (1500–2500 m) due to warmer temperatures and longer growing seasons [68,69]. Severe infestations not only reduce seed viability but also deplete soil seed banks, impairing natural regeneration [70]. This is particularly concerning given the slow growth rate of S. moorcroftiana. Similar pests in related leguminous species (e.g., Sophora flavescens) exhibit resistance to conventional pesticides, highlighting the need for species-specific management strategies [71].

4.1.3. Current Status of Diseases

Despite the scarcity of reported diseases in natural populations, S. moorcroftiana faces emerging disease risks under climate change and intensified cultivation. To date, no widespread fungal or bacterial pathogens have been documented in wild environments, likely due to the antimicrobial secondary metabolites produced by its endophytic microorganisms [32]. However, recent studies suggest potential vulnerabilities:
  • Powdery mildew (Erysiphales): During 2024 fieldwork in Xizang, powdery mildew infections were observed on greenhouse-cultivated S. moorcroftiana, manifesting as white mycelial growth on leaves and stems. This pathogen thrives under high humidity (>70%) and moderate temperatures (15–25 °C), conditions common in controlled environments [72]. Infected plants exhibited reduced photosynthetic capacity and stunted growth, highlighting risks for intensive cultivation systems.
  • Potential fungal threats: Comparative studies on related legumes (e.g., Sophora flavescens) indicate susceptibility to root rot (Fusarium spp.) and leaf spot (Alternaria spp.) under prolonged rainfall or poor drainage [73,74]. Although not yet reported in S. moorcroftiana, these pathogens could colonize stressed plants in waterlogged soils or monsoon-affected regions.
  • Bacterial risks: In arid regions, Xanthomonas spp. have caused stem cankers in drought-stressed leguminous shrubs [75]. Such pathogens may exploit physiological weaknesses in S. moorcroftiana during extreme drought heatwave events.
These observations underscore the need for proactive disease surveillance, particularly under shifting climatic conditions and expanded cultivation efforts.

4.2. Control Strategies

Pests primarily damage the pods and seeds of S. moorcroftiana during its developmental stage from June to July [76]. Due to the cryptic nature of seed pests, effective prevention and control remain challenging, making them a central focus of research efforts. Studies utilizing artificial caging and quantitative pest inoculation experiments have explored the relationship between pest density and seed yield loss. A significantly positive correlation was seen between pest infestation rates and yield reductions as pest density increases [77]. Moreover, research has indicated that pod and seed damage caused by surface arthropods is more pronounced in low-altitude S. moorcroftiana shrublands, particularly within certain altitudinal ranges [78]. These results highlight the importance of understanding the ecological and environmental factors influencing pest dynamics to develop effective, targeted pest management strategies. To control E. zinckenella and B. sophorae, an integrated pest management (IPM) approach is recommended, combining manual intervention, chemical control, and ecological management.

4.2.1. Control of Etiella zinckenella

  • Chemical control: A 5% chlorantraniliprole suspension concentrate is highly effective against E. zinckenella. When applied 3 and 7 days after fertilization, this pesticide achieved 100% efficacy in controlling larvae that damage flowers and 98.26% and 94.91% efficacy in controlling larvae infesting pods, respectively [79]. Additionally, a 25% spinetoram water-dispersible granule was found to be effective, although slightly less potent than chlorantraniliprole.
  • Ecological control: Emamectin benzoate and fresh neem oil significantly reduce pod borer populations while having minimal impact on parasitoid species. These products are particularly recommended when pest control must be balanced with ecological conservation [80].

4.2.2. Control of Bruchophagus onois

  • Plant quarantine: Strict regulations prohibit the transport of infested seeds to pest-free areas. Rigorous quarantine protocols must be enforced during seed transportation to prevent pest spread.
  • Seed treatment: Seed flotation in water can help remove damaged seeds, while fumigation with aluminum phosphide effectively eliminates pests from infested seeds.
  • Forest management: Severely infested plantations may require coppicing to remove infected plants, disrupt the habitat of B. onois, and reduce pest sources.
  • Chemical control (B. onois): During the adult emergence period of B. onois, two applications of a 50% malathion emulsifiable concentrate diluted 1:1000 are recommended. The first application should occur from late June to early July to target first-generation adults, and the second from late August to early September to control the second-generation adults [81].

4.2.3. Recommendations for Integrated Pest Management (IPM)

  • Ecological impact mitigation: Minimize non-target effects on pollinators and natural enemies by using selective pesticides (e.g., emamectin benzoate) and habitat management [82].
  • Long-term monitoring: Establish regional pest surveillance networks to track population dynamics and predict outbreaks [83].
  • Climate-adaptive disease monitoring: Establish disease risk models incorporating regional climatic variables (e.g., humidity and precipitation) to predict outbreaks of powdery mildew and other humidity-dependent pathogens [84].
  • Resistant germplasm screening: Screen wild S. moorcroftiana populations for disease-resistant traits, particularly against powdery mildew and root rot, to inform breeding programs [85].
  • Endophyte-based biocontrol: Leverage antimicrobial endophytes isolated from S. moorcroftiana seeds (e.g., Bacillus subtilis strains) to develop biofungicides targeting emerging pathogens [86].
  • Given the cryptic nature of E. zinckenella and B. onois, alongside their significant impact on S. moorcroftiana seed production, the strategy of “prevention first, integrated control” is strongly advised.
  • Early detection: Integrating molecular techniques with field monitoring can enable the precise prediction of pest occurrence and severity, enhancing early detection and intervention capabilities.
  • Rotational use of pesticides: To mitigate the development of resistance, it is crucial to avoid reliance on a single pesticide. Alternating between various highly effective, low-toxicity chemicals can disrupt pest adaptation and prolong the efficacy of pest control measures.
  • Eco-friendly strategies: Effective habitat management and the use of natural enemies, such as parasitoid wasps, can be optimized to regulate pest populations, offering a sustainable and environmentally friendly approach to pest control.
  • Disease research and control: In-depth studies on the antimicrobial mechanisms of endophytic microorganisms associated with S. moorcroftiana should be conducted to explore their potential in developing novel methods for preventing emerging plant diseases.

5. Conservation of Germplasm Resources of S. moorcroftiana

5.1. Current Population Distribution and Degradation Issues

The natural populations of S. moorcroftiana in Tibet are in a steady decline, with resources progressively diminishing. According to Luo Huaibin [87], these populations are predominantly found in arid desert areas regions, often forming monocultures where S. moorcroftiana is the dominant species. Populations on mobile sandy lands tend to be more evenly distributed and denser, whereas those on gravelly sand and sand dunes exhibit lower densities and greater instability, showing signs of severe degradation. In addition to natural factors, human activities and environmental pressures significantly contribute to the degradation. For example, local herders have long relied on S. moorcroftiana for fuel and livestock fodder, leading to a continuous decline in its natural population. The presence of insect pests further exacerbates this decline. Grazing disturbance also has a substantial impact on S. moorcroftiana communities. Under grazing conditions, the community formed by S. moorcroftiana and Chrysopogon aciculatus displays a richer species composition, higher diversity indices, and more robust community development [88].
The high demand for S. moorcroftiana’s alkaloids, particularly matrine and oxymatrine, has led to overharvesting in regions like the Yarlung Tsangpo River Basin. This overexploitation, combined with habitat destruction from grazing and land reclamation, has significantly reduced natural populations, especially in areas with high human activity. Intensive cultivation for medicinal purposes has also introduced new risks, such as disease outbreaks in controlled environments, further threatening wild populations.

5.2. Genetic Diversity

As a unique species native to the Yarlung Tsangpo River Basin, S. moorcroftiana plays a pivotal role in desertification control and ecological restoration. Current research suggests that the genetic diversity of S. moorcroftiana populations is generally low, particularly in the Linzhi region, where populations exhibit limited adaptability to environmental pressures, thus increasing the risk of species extinction [89]. Populations show significant genetic differentiation between high- and low-altitude areas, with gene flow being more frequent between populations in the middle reaches of the river and those in the upper and lower reaches. Wind-mediated seed dispersal is a key factor influencing this gene flow pattern [90]. Additionally, populations from different regions of S. moorcroftiana exhibit distinct genetic characteristics. In high-altitude areas, such as Shigatse, populations demonstrate higher genetic diversity, likely due to more favorable climatic conditions and lower levels of human disturbance. As climate change intensifies, areas suitable for S. moorcroftiana growth are expanding, with high-altitude populations becoming potential priority conservation areas [22]. Regular monitoring of genetic traits and gene flow, combined with an understanding of wind-driven seed dispersal, can enhance gene flow between populations, strengthening their adaptability and long-term survival prospects.

5.3. Conservation and Restoration Strategies

To address the degradation of S. moorcroftiana populations, a range of conservation and restoration strategies have been proposed. Restoration efforts should be tailored to the specific level of degradation in different areas. For severely degraded regions, artificial restoration techniques such as vegetation reconstruction and sand fixation technologies are recommended. In moderately degraded areas, measures such as artificial promotion and enclosure should be implemented to gradually restore community stability. For lightly degraded areas, rotational grazing and appropriate management practices can help maintain ecological balance and promote sustainable resource utilization [91,92].
Li Wenlong et al. [93] suggested that species diversity in S. moorcroftiana populations can improve under light grazing conditions. Disturbances, including grazing, enclosure, and flooding, significantly affect the community structure of S. moorcroftiana shrublands in the middle reaches of the Yarlung Tsangpo River [94]. Furthermore, climate change’s impact on S. moorcroftiana growth warrants close attention. While short-term increases in temperature may benefit growth, long-term temperature rises could limit its growth and lead to habitat shrinkage [95]. Therefore, bolstering the protection and artificial cultivation of S. moorcroftiana germplasm resources is essential not only for combating desertification and soil degradation on the Qinghai-Tibet Plateau but also for maintaining regional ecological balance.
Without effective conservation measures, S. moorcroftiana faces a high risk of local extinction in heavily harvested areas. However, ongoing efforts, such as germplasm conservation, habitat protection, and sustainable cultivation, offer hope for recovery. Future research should focus on optimizing cultivation techniques to meet medicinal demand while minimizing ecological impact. This includes breeding drought-resistant varieties, leveraging endophytic microorganisms for disease control, and promoting sustainable harvesting practices. By integrating medicinal use with ecological restoration, we can ensure the long-term survival of S. moorcroftiana while supporting its role in biodiversity conservation and ecological restoration.

6. Future Perspectives

6.1. Basic Research Directions

  • Deepening gene function studies: Future research should leverage gene-editing technologies such as CRISPR-Cas9 to validate the roles of these genes in drought tolerance and root development. Integrating transcriptomics and metabolomics will enable a comprehensive analysis of gene expression patterns under diverse environmental stresses, revealing molecular mechanisms that facilitate adaptation to plateau-specific adversities. Additionally, whole-genome resequencing of diverse germplasm resources will deepen the understanding of genetic diversity, population structure, and the genetic bases of adaptation, thereby supporting breeding and conservation initiatives.
  • Multi-omics integration: Combining genomic, transcriptomic, proteomics, and metabolomics data will allow the construction of a comprehensive molecular network model of drought adaptation.
  • Population genetics analysis: Resequencing geographically distinct populations will provide insights into genetic diversity and the adaptive evolution, supporting breeding and conservation initiatives. Specifically, investigating whether the expansion of sucrose metabolism-related genes is associated with evolutionary adaptations to arid environments will provide valuable insights and high-quality germplasm resources for genetic improvement.

6.2. Applied Research Directions

  • Breeding drought-resistant varieties: Molecular marker-assisted selection (MAS) and gene-editing technologies can expedite the development of drought-resistant S. moorcroftiana varieties.
  • Developing functional microbial agents: Probiotic microbial strains, such as nitrogen-fixing and phosphate-solubilizing bacteria, that promote root development and improve drought tolerance.
  • Exploiting functional compounds: Secondary metabolites in sucrose metabolism-related pathways should be further investigated for their roles in drought resistance and antioxidative properties.

6.3. Sustainable Utilization and Management

  • Habitat protection strategies: Strengthening environmental monitoring and establishing conservation zones to mitigate human-induced pressures on S. moorcroftiana populations.
  • Desertification control and ecological restoration: Harnessing S. moorcroftiana’s robust root systems and stress tolerance can enhance ecosystem stability in plateau and desertified regions.
  • Socioeconomic value promotion: Developing industries around S. moorcroftiana products, such as functional foods and medicinal products, can stimulate economic growth and conservation efforts.

7. Conclusions

In conclusion, S. moorcroftiana is a unique and ecologically significant species native to the Tibetan Plateau, with remarkable adaptability to extreme environments. Its secondary metabolites, particularly alkaloids like matrine and oxymatrine, hold significant medicinal potential, while its ecological functions in windbreak, sand stabilization, and soil conservation are critical for combating desertification and maintaining biodiversity. However, the species faces severe threats from overharvesting, habitat degradation, and climate change, necessitating urgent conservation efforts. This review synthesizes the latest research on S. moorcroftiana, providing a foundation for future scientific exploration, sustainable utilization, and conservation strategies.

Supplementary Materials

The following supporting information can be downloaded at https://www.mdpi.com/article/10.3390/f16030410/s1, Figure S1: Active compounds and clinical applications of S. moorcroftiana.

Author Contributions

Conceptualization and methodology, D.M. and S.Y. (Shuangyuan Yu); acquisition of data, D.M. and S.Y. (Sinong Yu); writing—original draft, D.M. and S.Y. (Sinong Yu); investigation, D.M., G.W., S.Y. (Shuangyuan Yu) and T.D.; writing—review and editing, D.M., S.Y. (Sinong Yu), F.C., S.Y. (Shuangyuan Yu), G.W. and T.D.; supervision, S.Y. (Sinong Yu), F.C., G.W. and T.D.; funding acquisition, F.C. and T.D. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the National Natural Science Foundation of China, grant number: 32471873; The STI 2030-Major Projects, grant number: 2023ZD0405605; the Natural Science Foundation of Jiangsu Province, China, grant number: BK20231291; and the China postdoctoral Science Foundation project, grant number: 2024M751426.

Data Availability Statement

Data is contained within the article or Supplementary Materials.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. Morphological characteristics of S. moorcroftiana in XiZang: (A) flower; (B) pod; (C) seed; and (D) seedling.
Figure 1. Morphological characteristics of S. moorcroftiana in XiZang: (A) flower; (B) pod; (C) seed; and (D) seedling.
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Figure 2. Resource distribution map of S. moorcroftiana in the Xizang region. The colors in the map range from light to dark, indicating the distribution area of the species in this distribution region. The lighter the color, the smaller the distribution area of the species in the corresponding region; the darker the color, the larger the distribution area of the species in that region.
Figure 2. Resource distribution map of S. moorcroftiana in the Xizang region. The colors in the map range from light to dark, indicating the distribution area of the species in this distribution region. The lighter the color, the smaller the distribution area of the species in the corresponding region; the darker the color, the larger the distribution area of the species in that region.
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Figure 3. Major pests (Bruchophagus onois) and diseases (Erysiphales) of S. moorcroftiana. (A) Seeds infested by the pest Bruchophagus onois, showing visible damage and deformities. (B) S. moorcroftiana plant affected by Erysiphales disease, demonstrating symptoms such as powdery mildew on the leaves.
Figure 3. Major pests (Bruchophagus onois) and diseases (Erysiphales) of S. moorcroftiana. (A) Seeds infested by the pest Bruchophagus onois, showing visible damage and deformities. (B) S. moorcroftiana plant affected by Erysiphales disease, demonstrating symptoms such as powdery mildew on the leaves.
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MDPI and ACS Style

Mei, D.; Yu, S.; Yu, S.; Cao, F.; Wang, G.; Dai, T. Unlocking the Potential of Sophora moorcroftiana (Fabaceae): The Overlooked Xizang Endemic. Forests 2025, 16, 410. https://doi.org/10.3390/f16030410

AMA Style

Mei D, Yu S, Yu S, Cao F, Wang G, Dai T. Unlocking the Potential of Sophora moorcroftiana (Fabaceae): The Overlooked Xizang Endemic. Forests. 2025; 16(3):410. https://doi.org/10.3390/f16030410

Chicago/Turabian Style

Mei, Duozhuoga, Sinong Yu, Shuangyuan Yu, Fuliang Cao, Guibin Wang, and Tingting Dai. 2025. "Unlocking the Potential of Sophora moorcroftiana (Fabaceae): The Overlooked Xizang Endemic" Forests 16, no. 3: 410. https://doi.org/10.3390/f16030410

APA Style

Mei, D., Yu, S., Yu, S., Cao, F., Wang, G., & Dai, T. (2025). Unlocking the Potential of Sophora moorcroftiana (Fabaceae): The Overlooked Xizang Endemic. Forests, 16(3), 410. https://doi.org/10.3390/f16030410

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