Research Advances in Chili Pepper Anthracnose

Tianjiao Sun

1,a  Affiliation:
1 College of Horticulture Hunan Agricultural University, Changsha, Hunan, 410125, China
All authors’ email:
a Email: 18390924928@163.com

All authors’ bio:
Tianjiao Sun was born in Turpan, Xinjiang. China, in 1995. She obtained a master’s
degree from Hunan Agricultural University. She currently studying at the HorticultureCollege of Hunan Agricultural University. His main research direction is ElucidatingMechanisms of Anthracnose Resistance and Innovating Germplasm in Pepper.

Abstract:

Anthracnose in chili pepper, caused by pathogenic fungi of the genus
Colletotrichum, primarily affects pepper fruits and leaves. It has become a significant
fungal disease in chili pepper production, causing global yield losses of 15%–50%and posing serious constraints to industry development. Currently, the complexspecies diversity of Colletotrichum pathogens and incomplete elucidation of keypathogenic mechanisms have limited the development of efficient control
strategies.This review summarizes research advances in pathogen identificationanddiversity, infection cycle and pathogenic mechanisms, host resistance and geneticbreeding, disease epidemiology, and integrated management techniques includingchemical and biological control. Future research directions are also discussed, aimingto provide insights for in-depth studies and effective management this disease. Keywords: Anthracnose in chili pepper; Colletotrichum; Pathogen diversity;Disease resistance genes; Integrated management

Introduction
Chili pepper anthracnose, caused by fungi of the genus Colletotrichumspp., is
one of the globally destructive diseases, often resulting in yield losses of 15%–50%[1]. According to the Food and Agriculture Organization of the United Nations (FAO)
statistics, China’s chili pepper cultivation area reached 18,857 ha in 2023, with a total
production of approximately 33.909 million tons
[2]. In recent years, the northwardshift of China’s rainfall belt has intensified anthracnose damage during the fruit
ripening stage in major producing provinces like Henan, Shandong, and Hebei, prompting a migration of the dry and processing pepper industries to more
climatically suitable regions
[3]. Traditional classification methods relyingonmorphology and single-gene ITS markers are prone to species misidentification. Furthermore, the complex species diversity of the pathogens and incompleteelucidation of key pathogenic mechanisms have constrained the development of
efficient control strategies to some extent. This review systematically summarizes thelatest research advances in chili pepper anthracnose, covering pathogen identificationand diversity, infection cycle and pathogenic mechanisms, host resistance and geneticbreeding, disease epidemiology, and integrated management techniques includingchemical and biological control. Future research directions are also discussed, aimingto provide a scientific basis for in-depth studies and effective management of this
disease. 1 Pathogen Diversity of Chili Pepper Anthracnose
1.1 Global Pathogen Complex of Chili Pepper Anthracnose
The classification system of the genus Colletotrichum underwent systematicrevision starting in 2009 based on multi-gene phylogenetic analyses. Integratingsequences of genes such as ITS, GAPDH, CHS-1, ACT, TUB2, CAL, SOD2, andAPN2, along with morphological characteristics, significantly improved species
identification accuracy
[4]. Currently, 24 species pathogenic to chili pepper have beenidentified globally, with 15 species reported in China
[5]. The core pathogencomplexes include: Colletotrichum scovillei (previously identified as C. acutatum);
Colletotrichum truncatum (previously identified as C. capsici); Colletotrichumsiamense (previously identified as C. gloeosporioides). Based on comprehensiveliterature reports, the global distribution of these major pathogens is summarizedinTable 1.Table 1: Global distribution of major Colletotrichum spp. causing chili pepper
anthracnose
Colletotrichum
Species Countries References C. truncatum
Australia, India, China, South Korea, Malaysia, USA, Vietnam, Thailand, Nigeria, Pakistan, Indonesia
[6-19]
C. scovillei
China, Brazil, South Korea, Malaysia, Indonesia, Japan, Philippines, Vietnam, Thailand, India, USA
[8,9,10,12,13,16,17,20,22,23,2
4,25,26,27,28,29,30]
C. siamense Australia, India, Brazil, Malaysia, Vietnam, Thailand, South
Korea, USA, China, Indonesia
[6,7,9,10,12,13,14,16,17,19,20, 23,27,29]
C. coccodes Brazil, Bulgaria, India, South Korea, Nigeria, USA, Vietnam [13,15,23,29,31,33]
1.2 Species Composition, Endemic Pathogens, and Novel Species Discoveries inChina
Systematic isolation and identification from diseased chili pepper samples across
29 provinces in China (yielding 1,285 isolates), combined with morphological andmulti-gene phylogenetic analyses (ITS/ACT/CAL/CHS-1/GAPDH/TUB2/HIS3), revealed 15 Colletotrichum species pathogenic to chili pepper in China. C. fioriniae, C. fructicola, C. siamense, C. scovillei, and C. truncatum were the most prevalent. Novel species first described include: C. conoides (Nanjing, Jiangsu), C. grossum(Haikou, Hainan), C. liaoningense (Xingcheng, Liaoning)
[9]. From rotten chili pepper
fruits in Sichuan Province, 88 isolates representing 7 Colletotrichumspecies wereobtained. Phylogenetic analysis based on morphology and multi-locus sequences
detected a novel species described as C. sichuanensis
[10]. In October 2021, approximately 60% of chili pepper fruits (cv. Lamb chili) in Yunnan Province, China, exhibited typical anthracnose symptoms. Multi-locus phylogenetic analysis distinctlydifferentiated isolate YN296-2 as C. jiangxiense, distinguishing it fromall other
species within the Colletotrichum species complexes
[36]. Morphological
characteristics of these endemic novel species are detailed in Table 2. Table 2: Taxonomic characteristics of novel Colletotrichum species endemic toChina
C. boninense Brazil, India [29,31]
C. dematium South Korea [23]
C. brevisporum Brazil, Vietnam, China [10,20,27,34]
C. fructicola India, China, Malaysia, Vietnam [7,10,12,27,29,35]
Colletotrichum
Species Region Colony
Appearance Growth
Rate （mm/day
）
Conidiophores Conidia Appressoria ReferencesLength × Width
（μm）
Shape Length × Width
（μm）
Shape Length × Width（μm）C. conoides Jiangsu Greyish white
aerial mycelium; reverse pale grey to medium grey,
13.25–13.7
5 (Mean
13.5)
22–30 × 3.5–5
Cylindrical
to clavate, both ends obtuse, granules evenly
13.0–17.5 × 5.0–6.5
（Mean
15.9×5.9）
Single or clustered, medium to dark
brown, ellipsoidal to
irregular, margins lobate
4.0–11.5 ×6.0–10.5（Mean8.35×7.1）[9]
2 Pathogen-Host Interaction
2.1 Disease Cycle and Pathogenesis
margin white distributed or deeply
incised
C. grossum Hainan White aerial mycelium; reverse pale grey, margin white
12.25–13.0
（Mean
12.6）
22–32 × 3–3.5
Cylindrical
to clavate, one end
acute, granules concentrate d at tip
14.5–20.5 × 5.0–7.5
（Mean
16.8 ×
6.3）
Single, medium
brown, ovoid to
irregular, margins lobate 5.5–11.5 ×4.0–10.5（Mean8.65 × 6.1 ）C. liaoningense Liaoning
Pale grey
aerial mycelium; reverse medium to
dark brown, margin white
12.0–12.75
（Mean
12.4）
27–30 ×
3.5–4.5
Cylindrical
to clavate, one end
acute, granules concentrate d at tip
14.0–18.5 × 5.0–7.5
（Mean
16.3 ×
6.1 ）
Single, medium
to dark brown, ellipsoidal to
irregular, margins lobate
3.5–5.0 ×2.5–4.5（Mean4.1 × 2.9）C. sichuanensis Sichuan
Initially white, maturing to
pale grey
aerial mycelium; reverse pale grey, sparse mycelium
6.1–6.4
（Mean
6.3 ± 0.1）
Branched
cylindrical, apex slightly
swollen Cylindrical, both ends obtuse
15–18.9 ×
5.4–6.5
（Mean
16.9 ×
6.2）
Light to dark
brown, irregular
to lobate 8.1–12.4 ×5.4–8.8（Mean10.2 × 6.8）[10]C. jiangxiense Yunnan
Dense cottony
aerial mycelium, white to grey, margin with
orange
conidial masses; reverse olive green with
pale orange margin
约 12.9 11.5–20 ×1–2.5
Cylindrical, ends obtuse or one acute
13–19 ×
4–6
（Mean
15.2 ×
5.2）
Light to dark
brown, circular/ellipsoi
dal/irregular
– [36]
Colletotrichum spp. can infect all parts of the chili pepper plant throughout its
growth cycle. During vegetative growth, leaves and stems are infected. Leaf
symptoms typically include circular or irregular necrotic lesions with greyish-white tolight brown centers and dark brown margins (Figure 1A, D). Severe infections causelesions to coalesce, leading to leaf death and abscission. Stem infections producebrown, sunken lesions or cankers (Figure 1B, E), which can girdle the stem, causingsystemic wilting of upper foliage. The most damaging phase occurs during fruit
ripening and post-harvest storage, particularly under high humidity conditions suchas
rainy seasons. Infected fruits initially develop small, water-soaked spots that rapidlyexpand into circular or nearly circular, deep brown to black, sunken necrotic lesions
(Figure 1C, F). This stage causes significant pre-harvest fruit drop and post-harvest
decay, leading to substantial yield and quality losses
[37, 38]. Figure 1. Characteristic symptoms of anthracnose on chili pepper fruit (A, D), leaves
(B, E), and stems (C, F)
A B CD E F
The core infection process of Colletotrichum spp. begins with conidiarecognizing hydrophobic signals on the host surface, germinating to formgermtubes
that differentiate into melanized appressoria at their tips. This structure accumulates
osmotic regulators like glycerol, generating enormous turgor pressure (5–8 MPa) todrive the mechanical penetration of the pepper cuticle by the penetration peg[39]. Notably, C. scovillei forms unique dendritic penetration pegs on chili pepper fruits –intermediate structures composed of thick-walled hyphal branches with sharp tips for
efficient cuticle penetration, a structure formed exclusively on chili fruit cuticles
[40].
Infection strategies subsequently diverge:Intramural necrotrophy (e.g., C. truncatum): Hyphae expand between the cuticle and cell wall, secreting toxins likecolletotrichin to induce rapid host cell necrosis (within 24 hours), bypassingabiotrophic phase
[41].Hemibiotrophy (e.g., C. siamense): After penetration, slender
primary hyphae develop and maintain a biotrophic phase inside cells for 3–5 days, during which effector proteins suppress host immunity. This phase transitions to thicksecondary necrotrophic hyphae that secrete pectinases and cellulases to destroy tissueand spread
[42]. 2.2 Key Pathogenicity-Related Genes
The pathogenicity of chili pepper anthracnose fungi is intricately regulated byamulti-gene network, involving infection structure development, nutritional strategytransition, and activation of key signaling pathways. Studies show that the NADPHoxidase complex encoded by CsNox2 and CsNoxR genes is crucial for penetrationpegformation. Knockout mutants (Δcsnox2, Δcsnoxr) form appressoria but fail topenetrate intact pepper epidermis, resulting in loss of pathogenicity
[43]. Pathogenesis
post-penetration heavily relies on conserved MAPKsignalingpathways: CsPMK1 (Fus3/Kss1 homolog) regulates appressorium formation and lipidmobilization via the homeobox transcription factor CsHOX7; its deletion causes
delayed conidial germination and complete loss of infectivity
[44]. CsSTE7 (MAPKK)
deletion leads to defects in appressorium formation and loss of pathogenicity [45]. Cell wall integrity pathway components CsMck1 (MAPKKK), CsMkk1 (MAPKK), and CsMps1 (MAPK) collectively regulate hyphal growth, stress response, andinfectious hypha development; mutants of all three lose pathogenicity
[46]. The adaptor
protein CsSte50, a key regulator of the MAPK cascade, is essential for pathogenicity;
its deletion causes impaired conidial germination, absence of appressoriumformation, and complete loss of virulence
[47]. 2.3 Host Resistance Mechanisms in Chili Pepper
Chili pepper resistance to anthracnose involves both preformed (passive) andinducible (active) defense mechanisms
[48]. Passive defense relies on pre-existingstructural barriers and antimicrobial compounds. Research indicates strong resistanceto non-wound inoculation, suggesting Colletotrichum cannot effectively penetrate theintact cuticle
[49]. In the resistant accession Capsicum baccatumPBC80, the fruit
epidermis provides initial defense via cuticle and cell walls. Post-infection, localizedcell wall thickening occurs, and high concentrations of reactive oxygen species (ROS)
and phenolic compounds directly inhibit pathogen expansion
[50]. Mature fruits exhibit
stronger resistance than immature fruits due to increased phenolic content (e.g., chlorogenic acid, caffeic acid) and reinforced cell wall structure
[51]. The CaAMP1protein strongly inhibits Colletotrichum spore germination and hyphal extensioninvitro
[52]. These barriers effectively hinder spore attachment, penetration, and initial
colonization. When pathogens breach passive defenses, chili pepper activates active defenseresponses via the innate immune system, centered on two cascading signals:
PAMP-triggered immunity (PTI) and effector-triggered immunity (ETI). PTI is
initiated when pathogen-associated molecular patterns (PAMPs) are recognizedbyplant pattern recognition receptors (PRRs), inducing MAPK signaling activation, Ca²⁺
influx, and ROS burst. This subsequently regulates defense gene expressionviasalicylic acid (SA) or jasmonic acid (JA)/ethylene (ET) pathways
[53]. ETI is triggeredwhen resistance proteins (e.g., NLR-type proteins) directly recognize pathogeneffectors, leading to a hypersensitive response (HR) involving programmed cell deathat infection sites to limit pathogen spread. The CbAR9 gene in C. baccatumPBC80encodes a CC-type NLR protein; its overexpression significantly enhances resistanceto anthracnose and activates pathogenesis-related (PR) gene expression(PR1, PR2, PR10), restricting pathogen diffusion
[54]. This process is finely regulatedby hormone signaling networks and transcription factors: SA signaling dominates
resistance to hemibiotrophic pathogens, activating PR genes and systemic acquiredresistance (SAR)
[55]. JA/ET pathways synergistically combat necrotrophic infections. Key transcription factors (WRKY, NAC, AP2-EREBP) act as regulatory hubs byintegrating hormone signals – e.g., WRKY33 positively regulates JA/ET signalingtopromote defensin PDF1.2 expression
[56], while CaWRKY50 negatively regulates SAsignaling
[55]. C2H2 zinc finger proteins (e.g., CanZF40/46) coordinate defense geneexpression like lipoxygenase (LOX) via JA/ET pathways [57]. Furthermore, activationof ripening-specific genes like PepEST (a JA-induced esterase) and thephenylpropanoid pathway (key genes 4CL, CCoAOMT) strengthens the multi-layeredsynergistic defense network in fruit resistance
[58, 59]. 3 Germplasm Resources Resistant to Chili Pepper Anthracnose
Resistance to chili pepper anthracnose exhibits significant species specificityandfruit developmental stage dependency. Commercial cultivars of C. annuumaregenerally susceptible, while broad-spectrum resistance primarily resides in wildrelatives C. baccatum and C. chinense
[60]. Several core germplasms withbroad-spectrum, high-level resistance have been identified (Table 3), showingsignificant pathogen specificity, fruit stage dependency, and genetic diversity. Table 3: Core resistant germplasms and their resistance characteristics
Colletotrichu m Species Resistant
Accession Pepper Species Resis
tance Level
Key Genetic Mechanism
Resistance Stability
Validation Population Referenc
es C. scovillei
PBC80 C. baccatum HR
Green fruit: Recessive gene Co4; Red fruit: Dominant gene Co5
(independently inherited)
Immunity
throughout
growth stages F2, BC1
[61]
PBC81 C. baccatum HR Polygenic control Broad-spectrum F2, BC1
[62]
The representative C. baccatum accession PBC80 exhibits immunity throughout
the growth cycle against 13 strains of both C. scovillei and C. truncatum(genestacking: green fruit recessive co4 + red fruit dominant co5)
[61]. Thebroadest-spectrum accessions PBC81 and PI594137 confer stable resistance tomultiple pathogens via polygenic or single dominant gene control
[65, 68]. While C.
PBC932 C. chinense R
Green fruit: 2 dominant
genes; Red fruit: 2
recessive genes Stage-specific BC1
[63]
GBUEL104 C. annuum HR
Dominant effect > Additive effect
Green/Ripe fruit
dominant
F2, BC1, BC2
[64]
Pi594137 C. baccatum R Single dominant gene Broad-spectrum
(stronger than PBC81/932)
F2, BC1
[65]
0038-9155 C. annuum HR
Green fruit: 2
complementary dominant
genes; Red fruit: 2
recessive duplicate
epistatic genes Stage-independen
t inheritance F2, BC1, BC2
[66]
C. truncatum
PBC932 C. chinense HR
Single recessive gene col
(Green & Red fruit)
Zero lesion, complete
resistance F2, BC1
[67]
PBC81 C. baccatum HR Polygenic control Broad-spectrum F2, BC1
[68]
83–168 C. annuum MR Single dominant gene Delays infection F2, BC1
[69]
Punjab Lal C. annuum R Single dominant gene Multi-pathogen
resistance F1、F2、BC1
[70]
Daepoong-cho C. annuum HR
Single recessive gene
(allelic to PBC932 col)
Suppresses initial
infection F2、BCR、BCS
[71]
C. siamense PRI95030 C. chinense HR
Major QTL B1 (Explains 57.3% variation)
Broad-spectrum F2
[19]
UENF1718/U
ENF1797
C. baccatum HR Stage-specific genes Symptom-free at all stages
–
[72]
baccatum shows strong resistance to C. truncatum and C. siamense, C. scovillei canpartially overcome resistance in the green fruit stage. The C. chinense accessionPBC932 achieves zero-lesion immunity to C. truncatum (single recessive col gene)
[67], but its resistance to C. scovillei is stage-specific (green fruit: two dominant genes;
red fruit: two recessive genes)
[63]. PRI95030 possesses resistance to both C. siamenseand C. truncatum via a major QTL B1 (explaining 57.3% lesion variation)
[19]. AmongC. annuum cultivars, a few highly resistant sources exist, such as GBUEL104 (Brazil)
[64], Daepoong-cho (Korea)
[71], and Punjab Lal (India)
[70], which achieve resistancevia dominant or recessive major genes but require interspecific introgressionfor
improvement. These resources combine resistance stability with agronomic value; for
instance, the hybrid DP-37×Putra chili 4 achieved synergistic improvement in diseaseresistance (lesion area <5%), yield (1273 g/plant), and pungency (31,236 SHU)
[73]. 4 Genetics and Mapping of Anthracnose Resistance in Chili Pepper
Genetic studies reveal that resistance to different pathogens is controlledbymajor genes, QTLs, or gene clusters, exhibiting significant pathogen species
specificity and fruit ripening stage dependency. Major resistance sources areconcentrated in wild species C. baccatum and C. chinense. Resistance introgressioninto cultivated C. annuum is accelerated through interspecific hybridizationandmarker-assisted selection (MAS). Reported molecular markers associated withanthracnose resistance traits for MAS are listed in Table 4. Table 4: Key QTLs/genes mapped for chili pepper anthracnose resistance
Colletotrichu m Species Resistant Accession QTL
Linkage Group/C
hromoso me Molecular Marker
(Type)
Marker Interval / Position References
C. scovillei
C. baccatum ‘PBC81’ CaR12.2 LG12 AFLP HpmsE032～EataMcgc01 [68]
C. baccatum ‘PBC81’ ARG_4a LG4 SNP chr4_161.7～chr4_207.43
[74]
ARG_4b LG4 SNP chr4_196.86
ARG_12 LG12 SNP chr12_36.95～chr12_39.89
ARR_11 LG11 SNP
chr11_232.44～chr11_234.14
C. baccatum ‘PBC80’ – LG12 SSR HpmsE032
[75]
C. chinense ‘PBC932’ – P5
SCAR/InD
el
SCAR-InDel
C.baccatum var. pendulum 881045
An9.1 P1 AFLP meD2emD3_2 [76]
C. baccatum ‘PBC80’
RA80rP2、RA80rP3.1、
RA80rHP1
LG4 SNP
BACSNP-4-63～BACSNP-4-60
RA80rP3.2 LG12 SNP [77]
BACSNP-12-61～BACSNP-12-58
RA80rHP2 LG9 SNP
BACSNP-3-84～BACSNP-3-87
C. chinense ‘PBC932’ AnRGO5、AnRGT5、
AnRGD5、AnRRO5、
AnRRT5
P5 InDel InDel～HpmsE116 [63]
AnRGO5 P5 KASPar P5L_P_81～P5L_P_137 [78]
AnRGO5 P5
KASPar、
InDel
P5in-2266-404～P5in-2267-978
[79]
C. baccatum ‘PBC80’ RA806_r1 P4 SNP SNP_305～331
RA806_g1 P8 SNP SNP_541～571 [80]
RA806_g2 P3 SNP SNP_228～218
6 个 C. chinense 种质：
IT229207、 IT236717、
IT261436、IT261448、
IT284059 、IT305437
– Ch01,、
Ch02,、
Ch03, Ch05、
Ch08、
Ch09
SNP
ChCh01: 37,429,407 bp
(CA01g29280)
Ch02: 146,421,570 bp
(CA02g17350)
Ch03: 1,918,096 bp
(CC-NBS-LRR)
Ch05: 未指定位置(PRP
基因)
[81]
C. baccatum ‘PBC80’ CcR9 LG9 AFLP HpmsE143～EtgtMcgt10 [54]
C. truncatum
C. annuum ‘Punjab Lal’ RCt1 –
ISSR、
AFLP
CtR-431 、CtR594
[70]
C. annuum ‘Punjab Lal’ QCoG-la.livr1.1 SSR HPMS725～CAMS644
For C. scovillei, wild species core accessions C. baccatum ‘PBC81’ and ‘PBC80’ contribute major QTLs CaR12.2 (chr12, AFLP markers HpmsE032~EataMcgc01→fine-mapped to a 36.95–39.89 Mbp NLR gene cluster)
[64,74] andRA80rP2/RA80rP3.1/RA80rHP1 (Chr4, SNP markers BACSNP-4-63~60)
[77], respectively. Resistance in ‘PBC80’ green fruit co-segregates completely with SSRmarker HpmsE032 (231 bp) (prediction accuracy 100%)
[75]. C. chinense accession’PBC932′ regulates resistance in both fruit stages via a QTL cluster at the endof
chromosome P5 (explaining 33.17–62.38% of green fruit resistance variation)
[63]. Its
key gene AnRGO5 (within a 164 kb interval, NBS-LRR protein CA05g17730) is
non-functional in susceptible materials due to structural variation
[79]. The linkedKASPar marker UN27353_1781 (2 cM) has a selection accuracy of 92.9%[78]. For C. truncatum, the CC-NLR gene CbAR9, cloned from C. baccatum ‘PBC80’ (locatedinmajor QTL CcR9, chr9) and functionally validated, activates PR gene pathways
[54]. The homologous locus in ‘PBC81’ has a SCAR marker with 82.5%selectionefficiency
[62]. The dominant gene RCt1 (resistance to C. truncatum) in C. annuum’Punjab Lal’ has a conserved STS marker CtR-431 (1.8 cM)
[70]. Significantly, breakthroughs have been made in understanding cross-resistanceQTL mechanisms: In C. baccatum ‘PBC81’, the major resistance locus for C. scovillei
(CaR12.2 on chr12) and the major locus for C. truncatum (CcR9 on chr9) areinherited independently. However, minor QTL CaR9 (chr9) and CcR12.2 (chr12)
co-localize with the major QTLs for the other pathogen
[62], revealing synergisticdefense pathways. Molecular marker pyramiding strategies developed based onthis
(e.g., combining SCAR-InDel and SSR-HpmsE032) increased resistant genotypeselection efficiency to 77.2%, significantly accelerating the introgression of resistancesources (e.g., PBC series) into cultivated types, providing theoretical and technical
support for multi-pathogen resistance pyramiding breeding
[75]. 5 Integrated Management of Chili Pepper Anthracnose
Integrated management of chili pepper anthracnose encompasses chemical control, development of microbial-based fungicides, utilization of plant-derived fungicides,
C. baccatum ‘PBC81’ CcR9 LG9 AFLP HpmsE143～EtgtMcgt03
[68]
CcRC LGC EaacMcgc02～EaatMccg07
C. chinense ‘PBC932’ RA932g、RA932r LG2 SNP
CAP_T39318_0_1_1042 ～CAP_ T22290_0_1_429
[77]
C. chinense ‘PBC932’ QTLUL P9 G31
[82]
QTL-L4 P11 G28
and breeding resistant varieties. The core framework of integrated strategies is
summarized in Figure 2. Figure 2: Management Strategies for Chili Pepper Anthracnose
5.1 Chemical Control
Chemical fungicide application remains crucial for controlling large-scaleepidemics before or during the initial outbreak of chili pepper anthracnose, due toits
rapid action, ease of application, and high efficiency. Through in vitro toxicity assays
and field trials, several highly effective fungicide classes have been identified, including picoxystrobin, thiophanate-methyl, difenoconazole, prochloraz, azoxystrobin, mancozeb, pyraclostrobin, and their combinations (e.g., difenoconazole·pyraclostrobin, fluazinam·pyraclostrobin)
[83-89]. However, prolongedsole or excessive use of chemicals is widespread, readily inducing pathogen resistanceand leading to significantly reduced efficacy
[88]. Furthermore, due to variations in field management practices, long-termsole or
excessive use is common, exacerbating the risk of pathogen resistance. Resistancestudies show significant differences in baseline sensitivity to the same fungicideamong different Colletotrichum species (e.g., C. truncatum vs C. scovillei). Fludioxonil exhibited the strongest inhibitory activity against these strains (EC50
values of 0.0120 μg/mL and 0.0142 μg/mL, respectively)
[90], while carbendazimanddifenoconazole also performed well. In Sichuan Province, C. scovillei showed highsensitivity to prochloraz (EC50 0.0337 μg/mL), tebuconazole, and epoxiconazole, providing a basis for regionalized precise chemical application
[91]. Regarding application technology optimization, chemical fungicides primarilywork by disrupting pathogen cell structures (e.g., increasing membrane permeability), interfering with metabolism, or inhibiting growth. Core measures include seedtreatment (e.g., copper sulfate, carbendazim) and field spraying
[92]. Recent studies
indicate significant synergistic effects from coformulations: 20%
fluazinam·pyraclostrobin SC (900 g/ha) achieved 82.65% control efficacy
[87]; 40%difenoconazole·pyraclostrobin SC and 25% pyraclostrobin SC were most effectivewhen applied at disease onset (88.59% and 86.99%, respectively)
[88]. Among singleagents, benzimidazoles (carbendazim, thiophanate-methyl), triazoles (hexaconazole, tebuconazole, epoxiconazole), and prochloraz-manganese chloride complexsignificantly reduced diseased fruit rates
[84]. Innovations in application strategies haveemerged: Using electric sprayers equipped with carbon spray booms and symmetricnozzles improved control efficacy by 4.90%~8.90%, saved 67.99%of spray solution, and reduced labor time by 55.51% [93]. 5.2 Microbial-based Fungicides
Facing environmental pressures, costs, and resistance risks associated withchemical control, microbial biocontrol agents (BCAs) represent an important greenstrategy for managing chili pepper anthracnose. Various beneficial microorganisms
exhibit significant antagonistic activity, with Bacillus spp. being particularlyprominent. Bacillus subtilis KB21 produces iturin-like cyclic lipopeptides (CLPs) that
combine direct inhibition of C. scovillei with induction of chili pepper defense geneexpression
[94]. Bacillus velezensis strains (e.g., LY7, HG-8-2, ZF438) performexceptionally: Endophyte LY7 enhances disease resistance and promotes growth; its
secreted Cupin-domain protein is a key antifungal factor
[95]. HG-8-2 andits
lipopeptide extracts show strong antagonism against C. scovillei
[96]. ZF438 achieved68.26% and 47.93% control efficacy on detached fruits and potted plants, respectively[97]. Bacillus subtilis subsp. endophyticus can colonize plants long-term, inhibitingpathogen growth and spore development via secreted peptide-like substances, potentially inducing plant defense enzymes
[98]. Bacillus amyloliquefaciens strains
achieved a 61.11% inhibition rate; their cell-free filtrates disrupted hyphae [99]. Bacillus subtilis JN032305 (on talc carrier) showed good stability and compatibilitywith carbendazim; seed treatment reduced disease incidence by 60%and significantlyincreased yield; root dipping increased root length by 120% and doubled yield; fruit
spraying reduced postharvest incidence by 65.3% [100]. Furthermore, filtrates andvolatile organic compounds (VOCs) from Brevibacillus sp. B-4359, specific strains of
Burkholderia spp., and some actinomycete isolates (8 isolates showed 100%inhibition) also demonstrated antifungal potential
[101-103]. The genus Trichoderma is a major fungal BCA. Trichoderma koningiopsis
PSU3-2 effectively controls postharvest anthracnose (C. siamense) through triplesynergistic mechanisms: nutrient competition and direct antagonism(79.57%inhibition), volatile organic compounds (VOCs, containing key components like2-phenylethanol, 38.33% inhibition), and mycoparasitism mediated by high-activitychitinase and β-1,3-glucanase secretion (causing hyphal deformation and lysis). Its
spore suspension (10⁶ spores/mL) as a dip treatment completely suppressed lesionformation (100% efficacy)
[104]. Li et al. screened 6 Trichoderma strains whosenon-volatile metabolites achieved over 92% inhibition; some volatile metabolites alsoshowed inhibitory activity
[105]. To enhance biocontrol efficacy, optimizationstrategies are being explored: A 3:7 ratio of B. velezensis LY7 (1.0×10⁸ CFU/mL)
with low-dose propiconazole (0.75 g/L) yielded optimal results
[106]; Radiationmutagenesis of B. velezensis successfully generated mutant strains with increasedinhibition rates (up to 59.9%)
[107]. 5.3 Plant-derived Fungicides
Developing natural and safe plant-derived fungicides is an important directionfor green control of chili pepper anthracnose. Extracts and active components fromvarious plants show significant antifungal effects. Effective Plant Extracts: Methanol
extracts of Justicia procumbens inhibited pathogen hyphal growth and sporedevelopment
[108]. Allicin (from garlic) performed well; its 6%aqueous emulsion(600~1200 μg/mL) achieved field spray control efficacy comparable to 50%carbendazim WP (1000 μg/mL)
[109]. 15% ginger extract also inhibited mycelial
growth and spore germination at levels comparable to synthetic fungicides
[110]. Plant
Essential Oils (EOs): Due to their broad-spectrum antifungal properties and natural
origin, EOs are a research hotspot. Litsea cubeba essential oil (LCEO) stronglyinhibited C. scovillei spore germination (MIC 0.60 mg mL⁻¹), primarily by disruptingcell membrane structure (causing spore deformation, content leakage, membrane lipidperoxidation) and inhibiting ergosterol synthesis
[111]. Clove oil (rich in eugenol) was
effective in vitro and for seed treatment: 0.25% concentration significantly reducedseed infection rate (from 75.8% to 7.5%) without affecting seed germinationor
seedling growth
[112]. Novel formulations are also being developed, suchas
oil-in-water nanoemulsions of thyme oil (TO) combined with licorice aqueous extract
(LAE)
[113]. 5.4 Breeding Resistant Varieties
Developing resistant cultivars is an economically efficient strategy to control
chili pepper anthracnose using genetic resistance, reducing pesticide reliance
[114]. Conventional breeding has successfully developed highly anthracnose-resistant
varieties like ‘UNEMAT Pedro’ (red fruit) and ‘UNEMAT Malagueta Pantaneira’ (orange-red hybrid fruit). Their fruit infection rates were significantly lower thancommercial cultivars, yields increased 2-4 times, and fruit quality was excellent
(distinct pungency, good storage)
[115]. CRISPR/Cas9 gene editing has achievedbreakthroughs in resistance breeding. By editing the susceptibility gene CaERF28, homozygous mutants free of foreign DNA were generated. These mutants (e.g., S1-14-1-9) showed significantly enhanced resistance, with disease indices droppingfrom 74.8% in wild type to <5% after fruit inoculation with Colletotrichum, whilemaintaining stable agronomic traits. This provides important material for developingnon-transgenic resistant chili pepper varieties
[116]. 6 Challenges and Future Perspectives
6.1 Current Challenges
Unclear Pathogen Population Structure: Although 15 Colletotrichumspecies
(including 5 novel species) have been identified in China, the population geneticstructure and dominant species in different production regions remain unknown. For
instance, the epidemiology of endemic pathogens like C. sichuanensis in SichuanandYunnan is not understood, hindering targeted breeding and precise chemical control
across regions. Insufficient Depth in Resistance Mechanisms: While Colletotrichuminfectionrelies on effector proteins to suppress host immunity, how these effectors specificallydisrupt plant defenses is not fully resolved. The molecular basis for the significant
resistance differences in the same cultivar at different fruit ripening stages alsoremains unclear, limiting the design of broad-spectrum resistant varieties. Difficulty in Transferring Superior Resistance Genes: Core resistance sources
like PBC80 and PBC932 are concentrated in wild species C. baccatumandC. chinense. Their introgression into cultivated C. annuum is hampered by reproductivebarriers and linkage drag. Although 12 major QTLs (e.g., CaR12.2, CcR9) have beenmapped, marker-assisted selection (MAS) efficiency is only 77.2%-92.9%, andpyramiding multiple genes entails long breeding cycles. Limited Field Efficacy of Biocontrol Agents: Microbial preparations andplant-derived active substances show high inhibition rates in vitro, but their fieldefficacy is often insufficient and inconsistent. Factors limiting large-scale applicationinclude inadequate colonization ability of biocontrol agents, instability of activeingredients, and unoptimized synergistic mechanisms with chemical fungicides. 6.2 Future Perspectives
Pathogen Surveillance and Characterization: Integrate multi-locus genotypingand population genomics to establish a national repository of Colletotrichumisolates
from major chili pepper production regions and develop a physiological racedifferentiation system. This will clarify the core pathogenic species in different
regions, providing a basis for deploying resistant varieties and implementing precisechemical control. Deciphering Effector-Host Interactions: Utilize yeast two-hybrid screeningandstructural biology to identify core Colletotrichum effector proteins and their cognatechili pepper immune receptors. Screen for corresponding resistance gene targets toprovide new directions for gene editing-based breeding. Advanced Resistance Breeding: Combine high-resolution genetic mapping withmulti-omics analysis to precisely map fruit ripening stage-specific resistance QTLs. Develop high-throughput KASP/InDel molecular markers. Overcome interspecifichybridization barriers using embryo rescue techniques. Employ gene editing to target
susceptibility genes. These approaches will enable efficient pyramiding of multipleresistance traits, significantly enhancing breeding efficiency. Optimized Integrated Management: Develop synergistic microbial-chemical
formulations to extend efficacy duration. Construct plant-derived synergistic systems
to enhance environmental stability. Implement precision deployment of resistant
varieties based on regional pathogen profiles. Establish a green management chain
from seed priming with beneficial microbes to postharvest preservation. References
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