1 Introduction

Hangjiao pepper is a common and important chili pepper variety in China and is widely grown in organic agriculture. Nowadays, many consumers prefer healthy food, so organic Hangjiao peppers that are free of pesticide residues and of good quality are very popular. However, in organic farming, problems caused by insufficient soil nutrients, continuous cropping and disease stress may be encountered, all of which can affect yield and quality (Gao et al., 2020; Zhang et al., 2024; Nagila et al., 2025). Therefore, to ensure the stable development of the Hangjiao pepper industry, it is necessary to improve the fertilization methods under organic conditions.

The way of fertilization will directly affect the growth, fruit quality and disease resistance of Hangjiao peppers. Studies have found that organic fertilizers such as amino acid water-soluble fertilizers can not only make Hangjiao peppers grow well and have a high yield, but also significantly increase components such as capsaicin, dihydrocapsaicin, protein and vitamin C (Bakpa et al., 2021). Combining organic fertilizer with straw, or using microbial organic fertilizer or bio-organic fertilizer, can also improve the types of microorganisms in the soil, increase soil enzyme activity, reduce soil-borne diseases, and enhance disease resistance (Gao et al., 2020; Liu et al., 2020; Zhang et al., 2024). In addition, choosing the appropriate rootstock for grafting and combining it with organic fertilizer management can also enhance the ability of Hangjiao peppers to cope with adverse environments (Nagila et al., 2025).

This study compared the effects of different organic fertilization methods on the pungency and disease resistance of Hangjiao pepper, and analyzed the mechanisms of action of multiple organic fertilizers and their combinations on growth, quality (especially spiciness) and disease resistance. This study aims to provide scientific basis and practical reference for achieving high yield, high quality and strong disease resistance of organic Hangjiao pepper.

2 Biological and Agronomic Basis of Pungency and Disease Resistance

2.1 Capsaicinoid biosynthesis pathways and influencing factors

Capsaicinoids are secondary metabolites unique to the Capsicum genus, mainly giving fruits a pungent taste (Feyera et al., 2024). Its synthesis involves two pathways: the phenylpropane pathway provides phenylalanine, the branched-chain fatty acid pathway provides precursors such as valine or leucine, and finally capsaicin is generated in the placenta tissue (Aza-González et al., 2011). There are some key enzymes in the synthesis process, such as AT3 (Pun1) and pAMT. pAMT is responsible for converting vanillin into vanillamine (Figure 1) (Kusaka et al., 2023; Nakaniwa et al., 2023). Capsaicin content is influenced by multiple factors, including genotype, placenta tissue specificity, fruit development stage, temperature, light, hormones and transcription factors (Arce-Rodriguez and Ochoa-Alejo, 2017; Burgos-Valencia et al., 2020; Sun et al., 2022). Some exogenous stimuli, such as chitosan and nano-selenium, can also increase the content of capsaicin by enhancing the expression of related genes (Kabita et al., 2020; Li et al., 2020).

Figure 1  Capsaicinoid production and the expression pattern of pAMT and its homologs in chili pepper (Adopted from Kusaka et al., 2023) Image caption: (a) Different tissues of chili pepper. IM: immature fruit stage 15 days after flowering (DAF), MG: mature green fruit stage 30 DAF, M: mature fruit stage 40 DAF. (b) Capsacinnoid accumulation in different tissues and expression levels of GABA-T homologs and Capsaicinoid biosynthesis-realated genes. All data are presented as means of three biological replicates. Error bars show standard errors (n = 3) (Adopted from Kusaka et al., 2023)

2.2 Mechanisms of disease resistance in Capsicum species

The disease resistance of chili peppers relies on multiple layers of defense, including physical barriers, R gene recognition, signal transduction (hormone pathways such as jasmonic acid, ethylene, abolic acid, etc.) and accumulation of secondary metabolites (Sun et al., 2019; Li et al., 2020). Capsaicin and its derivatives not only bring pungency, but also have antibacterial and antioxidant effects, which can directly inhibit pathogenic bacteria and improve the stress resistance of plants (Yasin et al., 2023). Some transcription factors (MYB, bHLH, AP2/ERF, etc.) regulate defense genes and metabolic pathways and participate in disease resistance responses (Liu et al., 2021; Wen et al., 2022; Islam et al., 2023; Song et al., 2023). Environmental stress and exogenous inducers (such as nano-selenium and chitosan) can also activate defense signals and enhance disease resistance (Kabita et al., 2020; Li et al., 2020).

2.3 Interactions between nutrient availability and secondary metabolite production

Nutrient supply (nitrogen, selenium, etc.) can affect photosynthesis, soluble sugar accumulation and hormone levels, thereby indirectly influencing the synthesis of capsaicin and other secondary metabolites. When nutrients are sufficient, the supply of precursor substances and energy is better, the activity of phenylpropane and fatty acid pathways is enhanced, and the contents of antioxidant substances such as capsaicin, flavonoids, and polyphenols will also increase. Nutritional status can also regulate hormone signals (such as jasmonic acid, ethylene), thereby affecting the expression of transcription factors and altering the types and contents of secondary metabolites (Sun et al., 2019; Li et al., 2020; Islam et al., 2023). In organic cultivation, reasonable fertilization management can optimize the quality of chili peppers and enhance their disease resistance.

3 Fertilization Regimes in Organic Hangjiao Pepper Production

3.1 Organic fertilization sources: compost, manure, green manure, and biofertilizers

Common organic fertilizers include compost, livestock and poultry manure (such as chicken manure), green manure and bio-fertilizers. Studies have found that the combined use of chicken manure and corn stalks can significantly increase the yield and disease resistance of Hangjiao pepper, improve the microbial structure in the soil, enhance soil functionality, and reduce continuous cropping obstacles and soil-borne diseases (Gao et al., 2020). Amino acid water-soluble fertilizer is a kind of bio-fertilizer. When used at a low concentration, it can increase the content of capsaicin and dihydrocapsaicin in the fruit of Hangjiao pepper, make the pungency stronger and the quality better (Bakpa et al., 2021).

3.2 Nutrient release dynamics and soil microbial interactions

The nutrients of organic fertilizer are released relatively slowly. It can provide crops with elements such as nitrogen, phosphorus and potassium for a long time and also promote the reproduction of beneficial microorganisms. The combined use of chicken manure and straw can not only increase the diversity of soil microorganisms, especially the dominant bacterial groups such as Proteobacteria and Ascomycota, but also enhance the disease resistance of the soil and reduce soil-borne diseases (Gao et al., 2020). Amino acid fertilizers can improve the physiological conditions and root activity of plants and help maintain the micro-ecological balance of the soil (Bakpa et al., 2021).

3.3 Comparative characteristics of balanced vs. targeted nutrient application

Balanced fertilization refers to the reasonable proportion of different nutrients to ensure the all-round growth of crops. Targeted fertilization is to adjust the nutrient ratio based on the specific needs of crops, such as increasing pungency or enhancing disease resistance. Studies have shown that low-concentration amino acid fertilizer (1.8 kg/ mu) can significantly increase capsaicin and protein content while ensuring yield, which helps to enhance pungency and quality (Bakpa et al., 2021). Chicken manure combined with straw is more suitable for improving soil quality and enhancing disease resistance, especially for plots with severe continuous cropping obstacles (Gao et al., 2020). However, if some fertilizers are used in excessive amounts, such as high-dose alkaline conditioning fertilizers, although the yield may increase, the benefits will gradually decrease (Ding, 2021).

4 Effects of Fertilization Regimes on Pungency

4.1 Influence of nitrogen, potassium, and microelements on capsaicinoid levels

The amounts of nitrogen, potassium and trace elements will directly affect the amount of capsaicin in Hangjiao peppers. Research has found that applying 1.8 kg per mu of amino acid water-soluble fertilizer can increase the capsaicin content of Hangjiao peppers to 5.80 mg/g and the dihydrocapsaicin content to 1.08 mg/g. At the same time, the levels of protein and vitamin C also increase, and the fruit quality is better. However, if the fertilizer concentration is too high, it does not necessarily lead to more capsaicin; instead, it may increase soluble sugar (Bakpa et al., 2021). This indicates that the dosage of nitrogen, potassium and trace elements should be reasonable in order to make the pungency of Hangjiao peppers better.

4.2 Timing and frequency of fertilization and its impact on pungency expression

The timing and frequency of fertilization are also very important. A study found that low-concentration amino acid fertilizer could promote capsaicin synthesis during the critical period of fruit growth by furrow application 6 weeks after transplanting (Bakpa et al., 2021). This suggests that for the organic cultivation of Hangjiao peppers, it is best to apply fertilizers in stages rather than all at once, as this makes it easier for the pungent substances to accumulate.

4.3 Environmental modifiers: soil type, moisture, and temperature under organic systems

In organic farming, soil type, moisture and temperature also affect capsaicin. They indirectly affect the pungency by altering the availability of nutrients and the types of microorganisms. Adding organic fertilizer and straw can increase the diversity of soil microorganisms, improve soil functions, and thereby enhance the stress resistance and quality of Hangjiao pepper (Gao et al., 2020). Under the action of organic fertilizer, capsaicin is more likely to accumulate stably when the soil maintains a neutral pH, moderate salinity and high organic matter (Xu et al., 2024). In addition, appropriate soil moisture and temperature can help with nutrient absorption and also make the enzymes related to capsaicin synthesis more active, thereby enhancing the pungency.

5 Effects of Fertilization Regimes on Disease Resistance

5.1 Role of nutrient balance in strengthening plant immune responses

The balance of nutrients is very important for plant immunity. Research has found that the appropriate use of amino acid water-soluble fertilizer can make the leaves of Hangjiao pepper greener, the root system more active and the protein content higher. These changes can enhance the basic immunity of plants and also improve their ability to cope with environmental stress (Bakpa et al., 2021). However, if too much nitrogen fertilizer is used, it may increase the number of plants with weak disease resistance in the field, thereby raising the risk of disease occurrence. This also indicates the importance of scientific fertilization and nutrient balance (Liu et al., 2017).

5.2 Impact on resistance to common pathogens in Hangjiao pepper

Combining organic fertilizer with straw can effectively reduce soil-borne diseases. For example, when chicken manure is mixed with corn stalks, it can not only increase the yield of Hangjiao pepper, but also improve the microbial community structure in the soil, making it more capable of resisting diseases such as anthracnose and bacterial wilt (Gao et al., 2020). This can alleviate the disease pressure caused by continuous cropping and make the Hangjiao peppers grow healthier.

5.3 Interaction of organic soil amendments with beneficial microbes for disease suppression

Organic fertilizers and bio-organic fertilizers can also promote the reproduction of beneficial microorganisms in the soil. These microorganisms can compete with pathogenic bacteria, inhibit each other, and induce plants to develop resistance, thereby reducing diseases (Gao et al., 2020). The disease-preventing strains in bio-organic fertilizers (such as Bacillus subtilis) can also cooperate with the beneficial bacteria in the soil to form a "microbial alliance", making Hangjiao pepper more resistant to diseases (Tao et al., 2020).

6 Integrated Analysis of Yield, Pungency, and Resistance

6.1 Trade-offs and synergies between high pungency and strong disease resistance

The pungency of chili peppers is mainly determined by the capsaicin content. Fertilization management can affect the pungency and disease resistance. Sometimes the two can restrict each other, or they may increase simultaneously. Studies have shown that appropriately reducing chemical fertilizers, or using organic fertilizers in combination with chemical fertilizers, can ensure yield while increasing capsaicin content and improving taste (Stan et al., 2021; Wang et al., 2022). Organic fertilizer combined with bio-fertilizer (such as mycorrhizal fungi and root-promoting bacteria) can not only increase yield and spiciness, but also improve the soil microbial environment, making plants more disease-resistant and stress-tolerant (Gao et al., 2020; Sini et al., 2024). Chicken manure combined with corn stalks can increase the diversity of soil microorganisms, reduce soil-borne diseases, and achieve a win-win situation of yield and disease resistance.

6.2 Multi-criteria assessment of fertilization regime performance

Different fertilization methods need to strike a balance among yield, pungency and disease resistance. 100% organic fertilizer, or a model with half organic fertilizer and half chemical fertilizer, can achieve high yield and quality in various types of chili peppers (Stan et al., 2021; Güneş et al., 2023). The combined use of bio-organic fertilizer, microbial agents and lime can not only increase yield and fruit quality, but also improve soil nutrients and microbial networks, making plants healthier (Gao et al., 2020; Sini et al., 2024; Zhang et al., 2024). Reducing the application of chemical fertilizers by 20% to 40% before harvest often does not lower the yield. Instead, it can increase capsaicin and flavor substances, and at the same time improve the efficiency of fertilizer utilization (Wang et al., 2022).

6.3 Potential for optimizing organic nutrient strategies for balanced quality and resilience

Optimizing organic nutrition management can simultaneously enhance the quality and stress resistance of Hangjiao pepper. The combined use of organic fertilizer, bio-fertilizer and straw can improve the structure of soil microorganisms, increase nutrient utilization rate and enhance plant health (Gao et al., 2020; Sini et al., 2024). Precise control of the application amounts of key elements such as nitrogen, phosphorus and potassium to avoid excess can stabilize the yield and improve capsaicin and disease resistance (Stan et al., 2021; Wang et al., 2022; Vadillo et al., 2024) In the future, multi-index optimization research should be strengthened to find the organic nutrient combination suitable for Hangjiao pepper, and achieve the organic cultivation goals of high yield, high quality and high resistance.

7 Case Study: Field Evaluation of Fertilization Regimes in Organic Hangjiao Pepper

7.1 Study location, soil conditions, and organic certification background

This case study was carried out in Lintao County, Gansu Province, China. It is a typical vegetarian-growing area and already has a foundation for organic agriculture certification. The experimental site was loam with medium fertility. The previous crop was also vegetables. Organic management has been carried out for many consecutive years without the use of chemical pesticides and chemical fertilizers, meeting the requirements of organic certification (Bakpa et al., 2021). Some experiments were also conducted on mountain and sandy soils to compare the effects of different soils on the growth and quality of Hangjiao pepper (Liu et al., 2020).

7.2 Experimental treatments, measurement of pungency, and disease resistance scoring

The experiment set up a variety of organic fertilization schemes, including amino acid water-soluble fertilizers of different concentrations (1.8, 2.7, 3.6 kg/mu), bio-organic fertilizers with Trichoderma guizhouense NJAU4742 as the core, organic fertilizers combined with corn stalks, and poultry manure combined with corn stalks. And combinations of microbial agents and lime, etc. (Liu et al., 2020; Zhang et al., 2024). The pungency was evaluated by the contents of capsaicin and dihydrocapsaicin in the fruit (Bakpa et al., 2021). Disease resistance is scored based on the incidence of field diseases and soil microbial diversity indicators, with a focus on the inhibitory effect of soil-borne diseases (Gao et al., 2020).

7.3 Results and farmer perspectives on practical applicability

The results showed that low-concentration amino acid water-soluble fertilizer (1.8 kg/ mu) could significantly increase the pungency of Hangjiao pepper (capsaicin content up to 5.80 mg/g), while increasing soluble protein, vitamin C and fruit quality (Bakpa et al., 2021). The combination of poultry manure and corn stalks not only increases yield, but also enhances soil microbial diversity and disease resistance, and reduces soil-borne diseases (Gao et al., 2020). Bio-organic fertilizer (Trichoderma guizhouense NJAU4742) can significantly increase yield (by 41.8% to 52.3%) under different soil types, and promote soil enzyme activity and nutrient transformation (Figure 2) (Liu et al., 2020). Farmers generally feedback that the scheme of low-concentration amino acid fertilizer and organic fertilizer combined with straw is convenient to operate, has moderate cost, and can also improve the quality and disease resistance of Hangjiao pepper. It is suitable for field promotion.

Figure 2  The pepper growth promotion in different treatments in sandy soil and mountain soil during two growth seasons (Adopted from Liu et al., 2020) Image caption: (A) The pot experiment results in sandy soil during the first season (FS); (B) the pot experiment results in mountain soil during the FS; C) the pot experiment results in sandy soil during the second season (SS); D) the pot experiment results in mountain soil during the SS; the treatments of the greenhouse pot experiment were as follows: T0: adding 0.01% (w/w, dw) chopped rice straw with the size of 1-2 mm; T1: 0.01% (w/w, dw) solid fermentation products; T3: 0.03% (w/w, dw) solid fermentation products; T6: 0.06% (w/w, dw) solid fermentation products; AA: 20% (v/w, dw) amino acid solution; AT1: 20% (v/w, dw) amino acid solution and 1% (w/w, dw) BOF (w/w, dw); AT3: 20% (v/w, dw) amino acid solution and 3% (w/w, dw) BOF; AT6: 20% (v/w, dw) amino acid solution and 6% (w/w, dw) BOF; BT: Bacillus amyloliquefaciens SQR-9-based BOF (stored in our lab); CK: chemical fertilizer (Adopted from Liu et al., 2020)

8 Challenges and Future Opportunities

8.1 Constraints in nutrient availability and consistency under organic farming

In organic farming, the stability of nutrient supply is a major issue. The nutrient release rate of organic fertilizers is slower than that of chemical fertilizers, and they are easily affected by environmental factors such as temperature and humidity. This will cause certain stages of nutrient deficiency in the growth of Hangjiao pepper, thereby affecting their pungency, yield and disease resistance. Research has found that water-soluble amino acid fertilizers of different concentrations can all improve the growth and quality of Hangjiao pepper, but the low-concentration treatment (1.8 kg) has the best effect in enhancing capsaicin and protein, etc. This indicates that the dosage and proportion of organic fertilizer need to be finely adjusted to achieve the best effect (Bakpa et al., 2021). Bio-organic fertilizers (such as those containing Trichoderma) can increase the yield and growth performance of Hangjiao pepper by promoting soil enzyme activity and nutrient transformation. However, its effect is affected by soil type and the rate of organic matter decomposition (Liu et al., 2020).

8.2 Research gaps in linking biochemical quality traits with disease resistance

At present, there are not many studies on the relationship between the pungency of Hangjiao pepper and its disease resistance. Although organic fertilizer combined with straw can enhance soil microbial diversity and increase disease resistance, the direct relationship mechanism between components such as capsaicin and disease resistance remains unclear. Most studies have focused on yield, quality improvement or changes in soil microorganisms. Systematic analyses of simultaneous changes in quality and disease resistance such as pungency and vitamin C are relatively lacking (Gao et al., 2020; Bakpa et al., 2021). In the future, multi-omics methods will be needed to reveal the connections between these qualities and disease resistance at the molecular and ecological levels.

8.3 Innovations in organic fertilization technology and integrated nutrient management

The new technology of organic fertilizer and comprehensive nutrient management are the key directions for improving the quality and disease resistance of Hangjiao pepper. Bio-organic fertilizers with the addition of specific microorganisms can regulate the soil microbial community, improve nutrient utilization efficiency and disease resistance (Liu et al., 2020). For example, organic fertilizers rich in Bacillus subtilis or Trichoderma can promote the proliferation of beneficial bacteria and form a synergistic disease-suppressing effect (Tao et al., 2020). In addition, the combined use of organic fertilizers with various organic substances such as straw and amino acids also helps to make the nutrient supply more stable and comprehensive (Gao et al., 2020; Liu et al., 2020). In the future, efforts should be focused on developing precisely formulated organic fertilizers, screening functional strains, and integrating intelligent nutrient management technology to achieve high-quality and high-disease-resistance goals in the organic cultivation of Hangjiao pepper.

Acknowledgments

The authors thank Dr. Xiao and Dr. Ma for their comments 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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