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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Microbiol.</journal-id>
<journal-title>Frontiers in Microbiology</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Microbiol.</abbrev-journal-title>
<issn pub-type="epub">1664-302X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fmicb.2024.1479957</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Microbiology</subject>
<subj-group>
<subject>Review</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>A comprehensive review of integrated management strategies for damping-off disease in chili</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes" equal-contrib="yes">
<name><surname>Delai</surname> <given-names>Chen</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>&#x002A;</sup></xref>
<xref ref-type="author-notes" rid="fn0001"><sup>&#x2020;</sup></xref>
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<contrib contrib-type="author" corresp="yes" equal-contrib="yes">
<name><surname>Muhae-Ud-Din</surname> <given-names>Ghulam</given-names></name>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>&#x002A;</sup></xref>
<xref ref-type="author-notes" rid="fn0001"><sup>&#x2020;</sup></xref>
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</contrib>
<contrib contrib-type="author">
<name><surname>Abid</surname> <given-names>Rimsha</given-names></name>
<xref ref-type="aff" rid="aff4"><sup>4</sup></xref>
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<contrib contrib-type="author">
<name><surname>Tian</surname> <given-names>Tian</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
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</contrib>
<contrib contrib-type="author">
<name><surname>Liu</surname> <given-names>Ruirui</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
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</contrib>
<contrib contrib-type="author">
<name><surname>Xiong</surname> <given-names>Yan</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
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</contrib>
<contrib contrib-type="author">
<name><surname>Ma</surname> <given-names>Shirong</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
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</contrib>
<contrib contrib-type="author">
<name><surname>Ghorbani</surname> <given-names>Abazar</given-names></name>
<xref ref-type="aff" rid="aff5"><sup>5</sup></xref>
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<aff id="aff1"><sup>1</sup><institution>School of Agriculture and Bioengineering, Longdong University</institution>, <addr-line>Qingyang</addr-line>, <country>China</country></aff>
<aff id="aff2"><sup>2</sup><institution>Gansu Key Laboratory of Protection and Utilization for Biological Resources and Ecological Restoration</institution>, <addr-line>Qingyang</addr-line>, <country>China</country></aff>
<aff id="aff3"><sup>3</sup><institution>Department of Plant Pathology, College of Agriculture, Guizhou University</institution>, <addr-line>Guiyang</addr-line>, <country>China</country></aff>
<aff id="aff4"><sup>4</sup><institution>Institute of Plant Protection, MNS University of Agriculture</institution>, <addr-line>Multan</addr-line>, <country>Pakistan</country></aff>
<aff id="aff5"><sup>5</sup><institution>National Key Laboratory of Green Pesticide, Key Laboratory of Green Pesticide and Agricultural Bioengineering, Ministry of Education, Center for R&#x0026;D of Fine Chemicals of Guizhou University</institution>, <addr-line>Guiyang</addr-line>, <country>China</country></aff>
<author-notes>
<fn fn-type="edited-by" id="fn0002">
<p>Edited by: Ravinder Kumar, Indian Agricultural Research Institute (ICAR), India</p>
</fn>
<fn fn-type="edited-by" id="fn0003">
<p>Reviewed by: Ashish Kumar Singh, ICAR-Vivekananda Parvatiya Krishi Anusandhan Sansthan, India</p>
<p>Abdul Gafur, SMF Corporate R&#x0026;D Advisory Board, Indonesia</p>
</fn>
<corresp id="c001">&#x002A;Correspondence: Chen Delai, <email>cdl829@126.com</email>; Ghulam Muhae-Ud-Din, <email>gm3085pp@outlook.com</email></corresp>
<fn fn-type="equal" id="fn0001">
<p><sup>&#x2020;</sup>These authors have contributed equally to this work</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>17</day>
<month>10</month>
<year>2024</year>
</pub-date>
<pub-date pub-type="collection">
<year>2024</year>
</pub-date>
<volume>15</volume>
<elocation-id>1479957</elocation-id>
<history>
<date date-type="received">
<day>13</day>
<month>08</month>
<year>2024</year>
</date>
<date date-type="accepted">
<day>30</day>
<month>09</month>
<year>2024</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#x00A9; 2024 Delai, Muhae-Ud-Din, Abid, Tian, Liu, Xiong, Ma and Ghorbani.</copyright-statement>
<copyright-year>2024</copyright-year>
<copyright-holder>Delai, Muhae-Ud-Din, Abid, Tian, Liu, Xiong, Ma and Ghorbani</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<p>Damping-off disease in chili (<italic>Capsicum annum</italic> L.) cultivation is a significant global issue, severely affecting seeds, seedlings, and young plants, regardless of the location of cultivation, whether in greenhouses or open fields. Despite chili being a widely popular vegetable used in various cuisines globally, farmers face challenges in meeting the growing demand due to the extensive damage caused by this disease, ranging from 20 to 85%. The shelf life and quality of mature pods are also severely affected. Damping-off disease is mainly caused by soil-borne fungus from the <italic>Pythium</italic> species, with additional contributions from <italic>Phytophthora</italic>, <italic>Fusarium</italic>, and <italic>Rhizoctonia</italic> species. These pathogens&#x2019; adaptability to diverse environmental conditions and resistance to synthetic fungicides make controlling damping-off on a commercial scale challenging. However, integrated disease management has shown promising results as a remedial approach. In this review, we discuss the current state of chili diseases, the nature of the pathogens causing damping-off, the epidemiology of the disease, and various control mechanisms. In this review, we broadly discuss the current state of chili diseases, the nature of the pathogens causing damping-off, the epidemiology of the disease, and various control mechanisms. Furthermore, we highlight the importance and efficacy of integrated disease management techniques, along with future prospects in unexplored areas, such as host&#x2013;pathogen interaction and sustainable disease control measures. The information in this review aims to assist chili growers in understanding the epidemiology and management of damping-off in chili cultivation.</p>
</abstract>
<kwd-group>
<kwd>biocontrol</kwd>
<kwd>chili cultivation</kwd>
<kwd>epidemiology</kwd>
<kwd><italic>Fusarium</italic> spp.</kwd>
<kwd><italic>Pythium</italic> spp.</kwd>
<kwd>resistant cultivars</kwd>
<kwd>sustainability</kwd>
</kwd-group>
<counts>
<fig-count count="1"/>
<table-count count="4"/>
<equation-count count="0"/>
<ref-count count="184"/>
<page-count count="17"/>
<word-count count="14441"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Microbe and Virus Interactions with Plants</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec sec-type="intro" id="sec1">
<label>1</label>
<title>Introduction</title>
<p>Chili (<italic>Capsicum annum</italic> L.) is a member of the Solanaceae family and is considered a vital vegetable <italic>cum</italic> spice, which is one of the most extensively cultivated crops worldwide. Morphologically, this is a heavily branched herbaceous annual shrub that grows well in sandy, fertile soil. The flowers are terminal, pentamerous, solitary, and bisexual, with a structure that allows the corolla to rotate. The stamens have blue to purple anthers attached at the base of the corolla (<xref ref-type="bibr" rid="ref11">Awang et al., 2013</xref>). Based on the fruit&#x2019;s shape, color, pungency, size, flavor, and use, chili is classified as a perennial crop and can be grown throughout the year. Approximately 400 different cultivars of chilies are grown, and well-developed pods are used as food ingredients for spices, sauces, pickles (<xref ref-type="bibr" rid="ref186">Zou and Zou, 2021</xref>), or beverages and medicine. In urban lifestyles, chili is grown as an ornamental plant in gardens. Chili contains a significant amount of minerals: potassium (K), iron (Fe), magnesium (Mg), and vitamins A and C, along with other significant nutrients (<xref ref-type="bibr" rid="ref146">Sarkar et al., 2022</xref>).</p>
<p>Chili originated in the tropical and subtropical regions of Central and South America and was introduced to Zhejiang Province (China) in 1591. It is now distributed across northern, western, and southern China (<xref ref-type="bibr" rid="ref185">Zou et al., 2020</xref>). China, Mexico, Indonesia, Turkey, and Spain are the leading green chili-producing countries, collectively responsible for approximately 75% of global production in 2013, according to <xref ref-type="bibr" rid="ref37">FAOSTAT (2020)</xref>. China, in particular, made the largest contribution, accounting for over half of the world&#x2019;s chili production, with a cultivation area exceeding 21,474&#x2009;km<sup>2</sup> in 2019 (<xref ref-type="bibr" rid="ref186">Zou and Zou, 2021</xref>). This surpasses the current land area allocated to versatile vegetables in the Chinese market. Moreover, more than 3,070&#x2009;km<sup>2</sup> is cultivated in Guizhou province (26.5982&#x00B0; N, 106.7072&#x00B0; E), which is higher than in any province in China.</p>
<p>Different crop genotypes are found in China, with productivity varying across provinces. The total chili yield in 2019 exceeded 64 million tons, which accounted for 7.76% of China&#x2019;s total national vegetable output. The total pepper yield in China reached 250 billion CNY, accounting for 11.36% of the total output value of national vegetables and contributing to 1.14% of the total farm income (<xref ref-type="bibr" rid="ref186">Zou and Zou, 2021</xref>). Though there is a rising demand for chili for versatile purposes, today, the supply chain has failed to cater to existing demand due to various challenges in chili cultivation. Among them, biotic stresses such as pest and disease attacks and abiotic stresses such as temperature fluctuations, water scarcity, and salinity are crucial. In addition, pre- and post-harvest losses due to poor management practices occurred on various scales.</p>
<p>Chili plant death in nurseries due to damping-off caused by pathogens has become a major issue in chili cultivation worldwide. The main causal organism is a soil-borne pathogenic fungus, <italic>Pythium</italic> spp., which affects chili plants in seedlings and younger roots. In addition, <italic>Rhizoctonia solani</italic>, <italic>Fusarium,</italic> and <italic>Phytophthora</italic> spp. are considered other pathogenic organisms that cause damping-off (<xref ref-type="bibr" rid="ref135">Rini and Sulochana, 2006</xref>). In this study, we have thoroughly discussed the damping-off disease in chili cultivation, current yield losses, sustainable crop management practices, integrated pest control methods, advancements in disease control technology, and its prospects.</p>
</sec>
<sec id="sec2">
<label>2</label>
<title>History and distribution of the host crop, <italic>Capsicum annuum</italic> L.</title>
<p><italic>Capsicum annuum</italic> L. has been known for over 9,500&#x2009;years. Chili is a native crop of Southern America and was first cultivated in Peru at approximately 7,500&#x2009;BC (<xref ref-type="bibr" rid="ref92">MacNeish, 1964</xref>). First, three species of chili, i.e., <italic>C. chinense</italic>, <italic>C. frutescens</italic>, and <italic>C. annuum,</italic> developed from common ancestors that grew widely in the north of the Amazon basin (NW-Brazil, Columbia), spreading to the other parts of America. Then, two more species, i.e., <italic>C. baccatum</italic> and <italic>C. pubescens,</italic> developed in different parts of America at the end of the eighteenth century (<xref ref-type="bibr" rid="ref62">IBPGR, 1983</xref>). The introduction of chili to China is endorsed by the journey of Columbus, who took the chili seeds from Spain, introduced them to Europe, and spread them to the subcontinents of Asia and Africa (<xref ref-type="bibr" rid="ref56">Heiser, 1976</xref>). Columbus confused the pungent fruits of <italic>Capsicum</italic> with black pepper <italic>Piper nigrum</italic> L., calling them red pepper owing to the red-colored fruits. <italic>Capsicum</italic> is not only related to the <italic>Piper</italic> genus. However, the terminology surrounding <italic>Capsicum</italic> is often confusing. Names such as chili, chili, aji, chile, pepper, and paprika denote the pungent fruits of <italic>Capsicum</italic>. The crop status spread quickly across Europe, moving to China, Japan, India, and Pakistan. Unlike other important cuisines, it became one of the most important cuisines in Asia and Europe (<xref ref-type="bibr" rid="ref186">Zou and Zou, 2021</xref>).</p>
<p>While there are reports of 20 wild species of chili peppers, only five species are currently used for cultivation: <italic>C. annuum, C. baccatum, C. frutescens, C. chinense, and C. pubescens.</italic> Among these, <italic>Capsicum annuum</italic> var. <italic>annum</italic> is the most extensively cultivated and economically significant among domesticated chili peppers. Notably, it is the only species commercially grown outside America, in Asia and Africa. Particularly in China, the crop was introduced to Zhejiang Province in 1591, and then it became the most important ingredient in various cuisines throughout the country (<xref ref-type="bibr" rid="ref186">Zou and Zou, 2021</xref>). The common name for chili in China is &#x201C;Lajiao&#x201D; in Chinese. The bell pepper is the non-pungent fruit of <italic>Capsicum</italic> and is known as &#x201C;Tianjin or Caijiao&#x201D; in Chinese. Over 95% of crops cultivated in China are <italic>C. frutescens</italic> L., <italic>C. annuum</italic> L., and <italic>C. chinense</italic> Jacq (<xref ref-type="bibr" rid="ref186">Zou and Zou, 2021</xref>), which are mostly cultivated in the northern and southern parts of China.</p>
</sec>
<sec id="sec3">
<label>3</label>
<title>Global production and Chinese contribution</title>
<p>Chili peppers are widely grown and consumed worldwide, with larger shares in Asia, Africa, America, and Europe. China is one of the major consumers, producers, and exporters of chili peppers (<xref ref-type="bibr" rid="ref159">Sundari et al., 2023</xref>). China&#x2019;s share of global chili pepper exports was approximately 13%. In 2020, global production was estimated at 37.9 million tons, with China accounting for approximately 36%. Mexico and Indonesia were the second and third largest green chili producers. According to the Guinness World Records and the Food and Agriculture Organization (FAO), China produces approximately 46% of the global chili supply to the market, as mentioned in <xref ref-type="table" rid="tab1">Table 1</xref> (<xref ref-type="bibr" rid="ref37">FAOSTAT, 2020</xref>; <xref ref-type="bibr" rid="ref139">Sachin et al., 2022</xref>).</p>
<table-wrap position="float" id="tab1">
<label>Table 1</label>
<caption>
<p>Annual chili production by the world&#x2019;s top producers in million tons (<xref ref-type="bibr" rid="ref37">FAOSTAT, 2020</xref>).</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="top">Country</th>
<th align="center" valign="top">Annual chili production in thousands of tons</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="top" colspan="2">Green chili</td>
</tr>
<tr>
<td align="left" valign="top">China</td>
<td align="center" valign="top">16,836</td>
</tr>
<tr>
<td align="left" valign="top">Mexico</td>
<td align="center" valign="top">2,818</td>
</tr>
<tr>
<td align="left" valign="top">Indonesia</td>
<td align="center" valign="top">2,773</td>
</tr>
<tr>
<td align="left" valign="top">Turkey</td>
<td align="center" valign="top">2,637</td>
</tr>
<tr>
<td align="left" valign="top">Spain</td>
<td align="center" valign="top">1,473</td>
</tr>
<tr>
<td align="left" valign="top">Egypt</td>
<td align="center" valign="top">867</td>
</tr>
<tr>
<td align="left" valign="top">Nigeria</td>
<td align="center" valign="top">767</td>
</tr>
<tr>
<td align="left" valign="top">Algeria</td>
<td align="center" valign="top">718</td>
</tr>
<tr>
<td align="left" valign="top">USA</td>
<td align="center" valign="top">566</td>
</tr>
<tr>
<td align="left" valign="top">Netherlands</td>
<td align="center" valign="top">430</td>
</tr>
<tr>
<td align="left" valign="top" colspan="2">Dry chili</td>
</tr>
<tr>
<td align="left" valign="top">India</td>
<td align="center" valign="top">1702</td>
</tr>
<tr>
<td align="left" valign="top">Thailand</td>
<td align="center" valign="top">322</td>
</tr>
<tr>
<td align="left" valign="top">China</td>
<td align="center" valign="top">308</td>
</tr>
<tr>
<td align="left" valign="top">Ethiopia</td>
<td align="center" valign="top">296</td>
</tr>
<tr>
<td align="left" valign="top">Bangladesh</td>
<td align="center" valign="top">158</td>
</tr>
<tr>
<td align="left" valign="top">Pakistan</td>
<td align="center" valign="top">141</td>
</tr>
<tr>
<td align="left" valign="top">Myanmar</td>
<td align="center" valign="top">140</td>
</tr>
</tbody>
</table>
</table-wrap>
<p>The growth in chili exports worldwide has been driven by production surpluses and the significant potential of the global chili trade. Since 2004, the volume of chili exports has experienced consistent growth, with an average annual growth rate of 6.04% (<xref ref-type="bibr" rid="ref153">Shimbov et al., 2019</xref>; <xref ref-type="bibr" rid="ref159">Sundari et al., 2023</xref>). However, interpreting global chili production trends and China&#x2019;s role in the industry is essential for policymakers, researchers, and stakeholders involved in the chili trade (<xref ref-type="bibr" rid="ref181">Yin et al., 2022</xref>).</p>
</sec>
<sec id="sec4">
<label>4</label>
<title>Uses and importance of chili</title>
<p>Chili is utilized in various forms, including fresh, green, or ripened fruits, along with dried and powdered forms. Fresh green pungent chili fruits are generally used in stuffing, salads, pickles, and as a flavoring agent in cooked meals, while fresh green non-pungent fruits are used as vegetables or processed with other food items for flavor. Additionally, highly pungent fruits are used as a spice to stimulate appetite and as a flavoring agent in ketchup. In addition to culinary uses, small quantities of chili are used in the cosmetic and garment industries (<xref ref-type="bibr" rid="ref148">Saxena et al., 2016</xref>).</p>
<p>Chilies have the potential to alleviate micronutrient deficiencies by supplying vitamins A, C, E, and B, along with minerals such as molybdenum, manganese, folate, potassium, and copper. They also supply significant macronutrients, such as protein, carbohydrates, fats, and dietary fiber, which are essential for human health (<xref ref-type="bibr" rid="ref125">Olatunji and Afolayan, 2018</xref>).</p>
<p>Peppers contain a wide range of phytochemicals, such as phenolics and flavonoids, which possess important antioxidant activities that help reduce the risk of degenerative diseases (<xref ref-type="bibr" rid="ref143">Saleh et al., 2018</xref>). Additionally, capsaicin in chili improves digestive health and nutrient absorption (<xref ref-type="bibr" rid="ref103">Marini et al., 2015</xref>; <xref ref-type="bibr" rid="ref137">Rosca et al., 2020</xref>), provides relief from joint pain (<xref ref-type="bibr" rid="ref38">Fattori et al., 2016</xref>), exhibits anti-inflammatory properties (<xref ref-type="bibr" rid="ref24">Chung and Campbell, 2016</xref>; <xref ref-type="bibr" rid="ref38">Fattori et al., 2016</xref>), and inhibits the growth of several types of fungi, including <italic>Candida albicans</italic> (<xref ref-type="bibr" rid="ref17">Behbehani et al., 2023</xref>).</p>
<p>Hot chili peppers also have antiviral properties, which can help prevent colds and flu while boosting the immune system (<xref ref-type="bibr" rid="ref105">Maurya and Sharma, 2023</xref>). Regular consumption of chili fruit is helpful against anorexia, varicose veins, hemorrhoids, and liver congestion. Chili extracts, both in pure and processed forms, are used externally as analgesic rubefacients for treating rheumatism, back pain, muscle pain, and articular and swollen feet, and are even used as antidotes for poisoning. Additionally, chilies have non-food and non-pharmacological uses, such as in the preparation of &#x201C;pepper sprays&#x201D; for self-defense and in making &#x201C;natural and organic pesticides.&#x201D;</p>
</sec>
<sec id="sec5">
<label>5</label>
<title>Chili plant morphology</title>
<p>The chili plant displays unique morphological features. The chili plant is a vigorously branched herbaceous species with primary, secondary, and tertiary branching. Its height typically ranges from 50 to 100&#x2009;cm (<xref ref-type="bibr" rid="ref127">Pawaskar, 2023</xref>). Its solitary, often paired, bisexual flowers have bell-shaped, twisted corollas with 5&#x2013;6 lobes. Notably, chili plants are primarily cross-pollinated by insects, with natural rates reaching up to 50% (<xref ref-type="bibr" rid="ref45">Gao et al., 2023</xref>). Chili peppers are distinctive berries with seeds not embedded in the pericarp (<xref ref-type="bibr" rid="ref12">Azlan et al., 2022</xref>). They change from green to red or purple upon ripening, with variations in shape, size, and pungency. Pungency is determined by capsaicin content and genetics, while the red color comes from capsanthin (<xref ref-type="bibr" rid="ref174">Wang et al., 2023</xref>). Orbicular seeds are developed within the fruit, and their size is influenced by nutrition (<xref ref-type="bibr" rid="ref16">Barboza et al., 2022</xref>). Chili plants, which are highly branched and have shallow roots, exhibit drought tolerance but struggle in waterlogged soil (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figure S1</xref>; <xref ref-type="bibr" rid="ref53">Guo et al., 2022</xref>).</p>
<sec id="sec6">
<label>5.1</label>
<title>Favorable climate for chili and chili diseases</title>
<p>Chilies thrive in tropical and subtropical regions up to 2000&#x2009;m in altitude, excluding pungent varieties (<xref ref-type="bibr" rid="ref1">Aashish et al., 2022</xref>). Optimal conditions include warm, humid climates that promote growth and fruit maturity, with an ideal annual rainfall of 850&#x2013;1,200&#x2009;mm. However, excessive rain and humidity can lead to poor fruit set and fruit rot. The ideal temperature for chilies ranges between 20&#x00B0;C and 25&#x00B0;C during the day and 15&#x00B0;C and 21&#x00B0;C at night (<xref ref-type="bibr" rid="ref7">Arom, 2022</xref>). A day length of 9&#x2013;10&#x2009;h stimulates plant growth, boosting productivity by 21&#x2013;24% and enhancing capsicum quality (<xref ref-type="bibr" rid="ref78">Kramchote and Suwor, 2022</xref>).</p>
<p>Soil is the main substrate for chili in open field conditions, and various other solid substrates, such as grow bags, are used under greenhouse conditions. Chili prefers well-drained, aerated soils rich in organic matter. Ideal soils are light loamy or sandy loamy with lime and organic content, while light soils require irrigation and organic fertilization. The 6&#x2013;7 pH range is ideal for optimum growth, and higher salinity hampers plant growth. Chili diseases have been a major reason for yield reductions in the world. Diseases caused by bacteria, viruses, fungi, and nematodes have badly affected chili crops worldwide (<xref ref-type="bibr" rid="ref157">Su et al., 2023</xref>; <xref ref-type="bibr" rid="ref178">Xie et al., 2023</xref>). Diseases associated with chili crops are summarized in <xref ref-type="table" rid="tab2">Table 2</xref>. One such fatal disease is damping-off, which occurs by soil-borne fungi genera, whether occurring pre- or post-plant emergence, accounting for up to 90% plant death and 62% seedling viability loss in nurseries and fields, as originally documented by <xref ref-type="bibr" rid="ref95">Majeed et al. (2018)</xref>.</p>
<table-wrap position="float" id="tab2">
<label>Table 2</label>
<caption>
<p>Biotic and abiotic stress of chili reported from different parts of the world.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="top">Disease name</th>
<th align="left" valign="top">Pathogen/abiotic stress</th>
<th align="left" valign="top">Symptom</th>
<th align="left" valign="top">Crop stage</th>
<th align="left" valign="top">Plant parts affected</th>
<th align="left" valign="top">Countries affected</th>
<th align="left" valign="top">Reference</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="top" colspan="7">Fungal diseases</td>
</tr>
<tr>
<td align="left" valign="top">Damping off</td>
<td align="left" valign="top"><italic>Pythium</italic> spp.<break/><italic>Fusarium</italic> spp.<break/><italic>Sclerotinia</italic> spp.</td>
<td align="left" valign="top">Failure of seedlings to emerge, soft, water-soaked, and discolored area at the base of the stem, stem constriction, seedling death, brown and rotted roots</td>
<td align="left" valign="top">seedling after transplantation</td>
<td align="left" valign="top">Roots and crowns of older plants</td>
<td align="left" valign="top">Worldwide</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref61">Hyder et al. (2021)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Black mold</td>
<td align="left" valign="top">
<italic>Alternaria alternata</italic>
</td>
<td align="left" valign="top">Rotting of fruits</td>
<td align="left" valign="top">Fruiting</td>
<td align="left" valign="top">Fruits</td>
<td align="left" valign="top">Tropical and sub-tropical countries</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref35">El-Garhy et al. (2020)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Phytophthora root rot</td>
<td align="left" valign="top">
<italic>Phytophthora capsici</italic>
</td>
<td align="left" valign="top">Rapid yellowing, wilting, soft rot, collapse of the rot, yellow-green needles, wilting, slow growth, dead branches, tree death</td>
<td align="left" valign="top">Seedling, fruiting</td>
<td align="left" valign="top">Roots</td>
<td align="left" valign="top">New Mexico</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref145">Sanogo and Carpenter (2006)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Verticillium wilt</td>
<td align="left" valign="top">
<italic>Verticillium dahliae</italic>
</td>
<td align="left" valign="top">Wilt, chlorosis, anthocyanescence, stunted and/or distorted growth, necrosis, and premature plant senescence.</td>
<td align="left" valign="top">Seedling</td>
<td align="left" valign="top">Leaves, stem, roots</td>
<td align="left" valign="top">New Mexico</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref144">Sanogo (2003)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Rhizoctonia root rot</td>
<td align="left" valign="top">
<italic>Rhizoctonia solaniis</italic>
</td>
<td align="left" valign="top">Stunting, yellowing of lower foliage, stem discoloration, reddish-brown dry cortical root rot, snapping off during high winds, reduced nodulation, yellowing or wilting of leaves</td>
<td align="left" valign="top">Seedling</td>
<td align="left" valign="top">Stem, roots</td>
<td align="left" valign="top">New Mexico</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref144">Sanogo (2003)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Frog eye leaf spot</td>
<td align="left" valign="top">
<italic>Cercospora capsici</italic>
</td>
<td align="left" valign="top">Circular lesions with a white canter, frog-eye, blighting of foliage, water-soaking of leaves, yellowing, reduced fruit size</td>
<td align="left" valign="top">Mature stage</td>
<td align="left" valign="top">Leaf</td>
<td align="left" valign="top">Taiwan</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref142">Salam et al. (2022)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Powdery mildew</td>
<td align="left" valign="top"><italic>Leveillula taurica</italic>, <italic>Oidiopsis taurica</italic></td>
<td align="left" valign="top">White, powdery, yellow, or chlorotic or powdery growth appearance on the upper leaf surface</td>
<td align="left" valign="top">Flowering and fruiting</td>
<td align="left" valign="top">Leaf</td>
<td align="left" valign="top">Places with a warm and dry climate</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref51">Glawe et al. (2010)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Anthracnose/ripe rot</td>
<td align="left" valign="top"><italic>Colletotrichum</italic> spp.</td>
<td align="left" valign="top">Leaf spots, fruit rot, stem cankers</td>
<td align="left" valign="top">Fruiting</td>
<td align="left" valign="top">Fruit</td>
<td align="left" valign="top">Tropical and sub-tropical countries</td>
<td align="left" valign="top"><xref ref-type="bibr" rid="ref164">Than et al. (2008)</xref> and <xref ref-type="bibr" rid="ref149">Saxena et al. (2014)</xref></td>
</tr>
<tr>
<td align="left" valign="top">Fruit rot</td>
<td align="left" valign="top">
<italic>Colletotrichum truncatum/capsici</italic>
</td>
<td align="left" valign="top">Sunken spots, collapse, discoloration</td>
<td align="left" valign="top">Mature plants</td>
<td align="left" valign="top">Fruit</td>
<td align="left" valign="top">Tropical and sub-tropical countries</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref148">Saxena et al. (2016)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Fusarium wilt</td>
<td align="left" valign="top"><italic>Fusarium solani</italic> var<italic>. capsici</italic></td>
<td align="left" valign="top">Yellowing and wilting, brown discoloration of the stem, and rotted roots</td>
<td align="left" valign="top">Seedling after Transplantation</td>
<td align="left" valign="top">Leaf</td>
<td align="left" valign="top">Leaf chlorosis, vascular discoloration, and wilting</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref160">Suryanto et al. (2010)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top" colspan="7">Bacterial diseases</td>
</tr>
<tr>
<td align="left" valign="top">Bacterial leaf spot</td>
<td align="left" valign="top"><italic>Xanthomonas campestris</italic> pv. <italic>vesicatoria</italic></td>
<td align="left" valign="top">Water-soaked spots, dark brown lesions, yellowing and dropping of leaves, deformation and twisting of young leaves</td>
<td align="left" valign="top">Seedling, flowering, fruiting</td>
<td align="left" valign="top">Leaf, stem, and fruits</td>
<td align="left" valign="top">Tropical and sub-tropical countries</td>
<td align="left" valign="top"><xref ref-type="bibr" rid="ref168">Vauterin et al. (2000)</xref> and <xref ref-type="bibr" rid="ref2">Abbasi et al. (2002)</xref></td>
</tr>
<tr>
<td align="left" valign="top">Bacterial soft rot</td>
<td align="left" valign="top"><italic>Erwinia carotovora</italic> pv<italic>. carotovora</italic></td>
<td align="left" valign="top">Watery lesions, tissue softening, sunken, water-soaked lesions, and wilting</td>
<td align="left" valign="top">Fruiting</td>
<td align="left" valign="top">Fruit</td>
<td align="left" valign="top">Areas with wet and cold climate conditions</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref156">Stommel et al. (1996)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Bacterial wilt</td>
<td align="left" valign="top">
<italic>Ralstonia solanacearum</italic>
</td>
<td align="left" valign="top">Yellowing, wilting, stunting, vascular necrosis, and vascular browning</td>
<td align="left" valign="top">After transplantation</td>
<td align="left" valign="top">Root and stem</td>
<td align="left" valign="top">Tropical and sub-tropical countries with high rainfall</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref123">Nguyen and Ranamukhaarachchi (2010)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Bacterial canker</td>
<td align="left" valign="top">
<italic>Corynebacterium michiganense</italic>
</td>
<td align="left" valign="top">Water-soaked margins and circular tan to dark spots on leaves, brown cankers on stems, and leaf curling,</td>
<td align="left" valign="top">Leaf and fruit</td>
<td align="left" valign="top">Leaves and stem</td>
<td align="left" valign="top">United States of America</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref86">Lewis Ivey and Miller (2000)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top" colspan="7">Viral diseases</td>
</tr>
<tr>
<td align="left" valign="top">Beet curly top virus</td>
<td align="left" valign="top">
<italic>Leafhopper transmitted Geminivirus</italic>
</td>
<td align="left" valign="top">Puckering, curling, yellowing, and stunting</td>
<td align="left" valign="top">Maturity</td>
<td align="left" valign="top">Whole plant</td>
<td align="left" valign="top">Western United States, Eastern Mediterranean basin</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref14">Bahari et al. (2022)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Pepper mottle virus</td>
<td align="left" valign="top">
<italic>Aphid transmitted Potyvirus</italic>
</td>
<td align="left" valign="top">Mild or bright yellow mottling on the leaves, leaf distortion, yellowing and stunting, blistering, and necrosis on fruit.</td>
<td align="left" valign="top">Maturity</td>
<td align="left" valign="top">Leaves and fruits</td>
<td align="left" valign="top">Florida, Southern USA, Arizona, Central America, Mexico, Thailand and India</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref71">Kaur et al. (2014)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Alfalfa mosaic virus</td>
<td align="left" valign="top"><italic>Aphid transmitted</italic> the <italic>Bacilliform virus</italic></td>
<td align="left" valign="top">Mottled leaves, irregular light and dark green areas on leaves</td>
<td align="left" valign="top">Maturity</td>
<td align="left" valign="top">Leaves</td>
<td align="left" valign="top">New Zealand</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref40">Fletcher (1983)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Pepper leaf curl virus</td>
<td align="left" valign="top">
<italic>Whitefly transmitted Geminivirus</italic>
</td>
<td align="left" valign="top">Upward curling of leaves, yellowing of veins, leaf size reduction, stunted growth, and yield reduction.</td>
<td align="left" valign="top">Maturity</td>
<td align="left" valign="top">Leaves and stem</td>
<td align="left" valign="top">United States, India, Nigeria, and South Asian countries</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref22">Chattopadhyay et al. (2008)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Pepper veinal mottle virus</td>
<td align="left" valign="top">
<italic>Aphid transmitted Potyvirus</italic>
</td>
<td align="left" valign="top">Stunting, curling, or twisting and yellow mosaic on the leaves, vein clearing, curling, twisting, and puckering of leaves and fruits.</td>
<td align="left" valign="top">Maturity</td>
<td align="left" valign="top">Leaves and fruits</td>
<td align="left" valign="top">Asian countries</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref114">Moury et al. (2005)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Pepper mosaic virus</td>
<td align="left" valign="top">
<italic>Aphid transmitted Potyvirus</italic>
</td>
<td align="left" valign="top">Mosaic and curling on leaves, fruit distortion, mottling symptoms on the leaf stem and fruits</td>
<td align="left" valign="top">Maturity</td>
<td align="left" valign="top">Leaf, stem, fruits</td>
<td align="left" valign="top">Argentina</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref3">Ahn et al. (2006)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Chili veinal mottle virus</td>
<td align="left" valign="top">
<italic>Aphid transmitted Potyvirus</italic>
</td>
<td align="left" valign="top">Dark green banding along veins on leaves, distorted leaves with mosaic patterns, twisted or fallen leaves, vein banding, reduced fruit size, severe puckering, thinning, shoestring, and cupping of leaf lamina in king chili plants, stunted growth with dark-green streaks on stems, mottled and deformed fruit, most flowers drop before fruit sets</td>
<td align="left" valign="top">Maturity</td>
<td align="left" valign="top">Leaf and fruits</td>
<td align="left" valign="top">Asian countries</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref114">Moury et al. (2005)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top" colspan="7">Nematode diseases</td>
</tr>
<tr>
<td align="left" valign="top">Root-knot nematode</td>
<td align="left" valign="top">
<italic>Meloidogyne incognita</italic>
</td>
<td align="left" valign="top">The prominent symptoms are of silting, yellowing, gall formation in roots, and stunted plants.</td>
<td align="left" valign="top">Maturity</td>
<td align="left" valign="top">Roots</td>
<td align="left" valign="top">Different parts of the world</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref165">Thiyagarajan (2014)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Stubby root nematode</td>
<td align="left" valign="top">
<italic>Paratrichadorus minor</italic>
</td>
<td align="left" valign="top">Stunted, chlorotic, wilted, and symptoms resemble nutrient deficiency.</td>
<td align="left" valign="top">At any stage</td>
<td align="left" valign="top">Roots</td>
<td align="left" valign="top">North America</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref87">Li et al. (2010)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Sting nematode</td>
<td align="left" valign="top">
<italic>Belonolaimus longicaudatus</italic>
</td>
<td align="left" valign="top">Stunted, chlorotic, wilted, and symptoms resemble nutrient deficiency</td>
<td align="left" valign="top">At any stage</td>
<td align="left" valign="top">Roots</td>
<td align="left" valign="top">North America</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref152">Shaver et al. (2017)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Root-lesion nematode</td>
<td align="left" valign="top">
<italic>Pratylenchus penetrans</italic>
</td>
<td align="left" valign="top">Stunted, chlorotic, wilted, and symptoms resemble nutrient deficiency, low flowering, and yield.</td>
<td align="left" valign="top">At any stage</td>
<td align="left" valign="top">Roots</td>
<td align="left" valign="top">North America</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref39">Figueiredo et al. (2021)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top" colspan="7">Insects and mite infection</td>
</tr>
<tr>
<td align="left" valign="top">Mite feeding injury</td>
<td align="left" valign="top">
<italic>Polyphagotarsonemus latus</italic>
</td>
<td align="left" valign="top">&#x201C;Inverted spoon-shaped&#x201D; leaves, pods with a rusty/corky surface</td>
<td align="left" valign="top">Seedlings and maturity</td>
<td align="left" valign="top">Leaves and fruit</td>
<td align="left" valign="top">Africa, Asia, Australia, South and North America, Pacific lands</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref169">Venzon et al. (2008)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Aphid feeding injury</td>
<td align="left" valign="top">
<italic>Myzus persicae Aphis gossypii</italic>
</td>
<td align="left" valign="top">Distorted, mottled young leaves, chlorosis, leaf drop, reduced fruit size</td>
<td align="left" valign="top">Seedlings and maturity</td>
<td align="left" valign="top">Leaves and fruit</td>
<td align="left" valign="top">India, Orient and Pacific lands, Sri Lanka, and Continental USA</td>
<td align="left" valign="top"><xref ref-type="bibr" rid="ref161">Tapia et al. (2008)</xref> and <xref ref-type="bibr" rid="ref167">Varghese and Mathew (2013)</xref></td>
</tr>
<tr>
<td align="left" valign="top">Thrips feeding injury</td>
<td align="left" valign="top">
<italic>Thrips parvispinus Scirtothrips dorsalis</italic>
</td>
<td align="left" valign="top">&#x201C;Boat-shaped&#x201D; curled leaves, distorted pods</td>
<td align="left" valign="top">Seedlings and maturity</td>
<td align="left" valign="top">Leaves and fruit</td>
<td align="left" valign="top">Worldwide</td>
<td align="left" valign="top"><xref ref-type="bibr" rid="ref94">Maharijaya et al. (2011)</xref> and <xref ref-type="bibr" rid="ref68">Johari et al. (2014)</xref></td>
</tr>
<tr>
<td align="left" valign="top" colspan="7">Abiotic stress</td>
</tr>
<tr>
<td align="left" valign="top">Blossom-end rot</td>
<td align="left" valign="top">Watering and calcium deficiency</td>
<td align="left" valign="top">Light green or yellow-colored sunken spot and water-soaked area that becomes a dark brown or black dry rot</td>
<td align="left" valign="top">At any stage</td>
<td align="left" valign="top">Fruit</td>
<td align="left" valign="top">Different parts of the world</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref102">Marcelis and Ho (1999)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Sunburn</td>
<td align="left" valign="top">Sun</td>
<td align="left" valign="top">White or black discoloration on fruits, brown or white foliage, leaves turning brown or ivory-white and becoming dry and crispy, yellow, bronze, or brown spots on the sun-exposed side of the fruit</td>
<td align="left" valign="top">At any stage</td>
<td align="left" valign="top">Fruit</td>
<td align="left" valign="top">Tropical and sub-tropical countries</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref89">L&#x00F3;pez-Mar&#x00ED;n et al. (2011)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Salt injury</td>
<td align="left" valign="top">Salt</td>
<td align="left" valign="top">Stunting or death of seedlings, reduction in leaf, flower, and fruit size, necrosis of leaf margins, less vegetative growth, reduced root growth</td>
<td align="left" valign="top">At any stage</td>
<td align="left" valign="top">Seedling</td>
<td align="left" valign="top">All over the world</td>
<td align="left" valign="top"><xref ref-type="bibr" rid="ref115">Mustafa et al. (2014)</xref> and <xref ref-type="bibr" rid="ref21">Butt et al. (2021)</xref></td>
</tr>
<tr>
<td align="left" valign="top">Wind injury</td>
<td align="left" valign="top">Wind</td>
<td align="left" valign="top">Wilting and death of young seedlings, broken stems, or branches</td>
<td align="left" valign="top">At any stage</td>
<td align="left" valign="top">Foliage</td>
<td align="left" valign="top">All over the world</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref128">Pohronezny et al. (1992)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Hail injury</td>
<td align="left" valign="top">Hail</td>
<td align="left" valign="top">Severe hail injury results in shredded leaves or defoliation-damaged stems and fruit and physical damage to the foliage</td>
<td align="left" valign="top">At any stage</td>
<td align="left" valign="top">Foliage and Fruit</td>
<td align="left" valign="top">All over the world</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref69">Joukhadar and Walker (2020)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Herbicide injury</td>
<td align="left" valign="top">Herbicide</td>
<td align="left" valign="top">Leaf cupping or curling, petiole twisting, epinasty, blotches or discoloration on the leaves, change of leaf color, severe tissue distortion or deformation</td>
<td align="left" valign="top">Maturity</td>
<td align="left" valign="top">All parts of the plant</td>
<td align="left" valign="top">All over the world</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref44">Galloway et al. (2000)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">Nutrient deficiencies and toxicities</td>
<td align="left" valign="top">Nutrients</td>
<td align="left" valign="top">Stunted growth, chlorosis, interveinal chlorosis, and purplish-red discoloration or premature dying of leaves</td>
<td align="left" valign="top">At any stage</td>
<td align="left" valign="top">All parts of plants</td>
<td align="left" valign="top">All over the world</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref73">Khaitov et al. (2019)</xref>
</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
</sec>
<sec id="sec7">
<label>6</label>
<title>Damping-off disease on chili</title>
<p>Damping-off is a significant fungal disease affecting chili production worldwide, leading to substantial economic losses. The disease primarily targets seedlings, causing poor germination and plant death, which reduces overall crop yield. Farmers experience financial setbacks due to the need for replanting, increased use of fungicides, and lower productivity. In regions highly dependent on chili cultivation, such as China, India, and Mexico, the disease poses a significant threat to growers&#x2019; livelihoods, disrupting both local and global markets. The chili crop is infected by more than 100 different types of plant pathogens during its vegetative and reproductive stages (<xref ref-type="bibr" rid="ref64">Jayapala et al., 2019</xref>). Damping-off disease is caused by soil-borne pathogens such as <italic>Pythium</italic> sp., <italic>Phytophthora</italic> sp.<italic>, Rhizoctonia solani, and Fusarium</italic> spp. (<xref ref-type="bibr" rid="ref162">Thakur and Singh, 2023</xref>).</p>
<p>Damping-off is one of the most critical diseases that leads to the decay of germinated seeds and young seedlings, which causes a huge economic loss for farmers in nurseries and fields. These fungal or fungal-like organisms cause seed death during germinating in nurseries, with losses of up to 90% under pathogen-favorable conditions (<xref ref-type="bibr" rid="ref9">Arora et al., 2021</xref>). Damping-off on chili occurs in two stages: the pre-emergence and post-emergence phases. In the pre-emergence phase, disease typically begins during the seedling stage, where the fungi invade seeds or young plants in waterlogged or poorly drained soil. Initial symptoms include seed rot or germination failure, leading to poor crop emergence. Infected seedlings may develop water-soaked lesions at the base of the stem, which gradually turn brown, causing the stem tissue to collapse. This pre-emergence damping-off stage is particularly destructive, as the seedlings die before breaking the soil surface (<xref ref-type="bibr" rid="ref95">Majeed et al., 2018</xref>). However, the stem tissue near the soil line becomes soft, sunken, and girdled during the post-emergence phase. The affected seedlings wilt, fall over, and die due to disrupted nutrient and water transport, a condition often referred to as &#x201C;wire stem&#x201D; (<xref ref-type="bibr" rid="ref83">Lamichhane et al., 2017</xref>).</p>
<p>This deterioration often results in reduced seedling vigor, and the seedlings may die near the coleoptile, exposing whitish fungal growth on the surface. In the initial stages, affected plants exhibit wilting, eventually leading to a severe infestation. Surviving plants display stunted growth, and the affected areas often exhibit irregular and uneven development (<xref ref-type="bibr" rid="ref175">Were et al., 2023</xref>). Damping-off symptoms can generally be observed from seeding until the 4th to 6th week post-sowing (<xref ref-type="supplementary-material" rid="SM2">Supplementary Figure S2</xref>). In this study, we focus mainly on <italic>Pythium</italic> spp. (<xref ref-type="bibr" rid="ref83">Lamichhane et al., 2017</xref>) regarding damping-off. Several species of <italic>Pythium</italic> associated with chili to cause damping-off disease in different parts of the world are mentioned in <xref ref-type="table" rid="tab3">Table 3</xref>.</p>
<table-wrap position="float" id="tab3">
<label>Table 3</label>
<caption>
<p>Fungal species associated with damping-off of chili in different parts of the world.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="top">Pathogen</th>
<th align="left" valign="top">Country/Province</th>
<th align="left" valign="top">Reference</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="top">
<italic>Pythium cedri</italic>
</td>
<td align="left" valign="top">Jiangsu/China</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref133">Rai et al. (2020)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>Fusarium oxysporum</italic>
</td>
<td align="left" valign="top">Guizhou/China</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref90">Luo and Yu (2020)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. helicoids</italic>
</td>
<td align="left" valign="top">Guangdong/China</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref23">Chen et al. (2016)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top"><italic>Pythium</italic> spp. <italic>Fusarium</italic> spp., <italic>Phytophthora</italic> spp.</td>
<td align="left" valign="top">Heilongjiang/China</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref28">Dai et al. (2013)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top"><italic>Pythium</italic> spp.</td>
<td align="left" valign="top">Hainan/China</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref61">Hyder et al. (2021)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top"><italic>P. breve</italic> and <italic>P</italic>. <italic>baisense</italic></td>
<td align="left" valign="top">South China</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref88">Long et al. (2012)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. myriotylum</italic>
</td>
<td align="left" valign="top">Pakistan</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref61">Hyder et al. (2021)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. aphanidermatum</italic>
</td>
<td align="left" valign="top">India</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref116">Muthukumar et al. (2010)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. graminicola</italic>
</td>
<td align="left" valign="top">India</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref31">Dubey et al. (2019)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. ultimum</italic>
</td>
<td align="left" valign="top">India</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref182">Zagade et al. (2016)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. spinosum</italic>
</td>
<td align="left" valign="top">Pakistan</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref122">Nawaz et al. (2016)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. diliense</italic>
</td>
<td align="left" valign="top">India</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref118">Muthukumar et al. (2018)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. heterothallicum</italic>
</td>
<td align="left" valign="top">India</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref118">Muthukumar et al. (2018)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. debaryanum</italic>
</td>
<td align="left" valign="top">Pakistan</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref151">Shahid et al. (2017)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. intermedium</italic>
</td>
<td align="left" valign="top">Pakistan</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref121">Nawaz et al. (2017)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. debaryanum</italic>
</td>
<td align="left" valign="top">India</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref52">Gomathi et al. (2012)</xref>
</td>
</tr>
</tbody>
</table>
</table-wrap>
<sec id="sec8">
<label>6.1</label>
<title>The biology of different pathogens causing damping-off</title>
<p><italic>Pythium</italic> species are a group of soil-borne pathogens known for causing damping-off and root rot in a wide range of plants, including chili. They thrive in wet, poorly drained soils and attack seedlings, leading to stunted growth and plant death. Some <italic>Phytophthora</italic> species can cause damping-off in seedlings. For instance, <italic>Phytophthora capsici</italic> is known to cause damping-off in seedlings of various crops, including peppers and other vegetables. The pathogen attacks young seedlings, leading to poor germination, stem collapse, and death, similar to other damping-off pathogens such as <italic>Pythium</italic> and <italic>Rhizoctonia</italic>. <italic>Rhizoctonia solani</italic> is a soil-borne fungal pathogen that affects chili seedlings by attacking their roots and stems, leading to poor germination, wilting, and eventual death of the young plants. <italic>Rhizoctonia</italic> thrives in warm, moist conditions and can lead to significant crop losses if not managed effectively. <italic>Fusarium</italic> species are soil-borne fungi that attack chili seedlings&#x2019; roots and lower stems, leading to poor emergence, wilting, and seedling death. <italic>Fusarium</italic> thrives in warm, moist soils and can cause significant damage if conditions are favorable.</p>
</sec>
<sec id="sec9">
<label>6.2</label>
<title>Epidemiology of the disease</title>
<p>Environmental factors play a key role in spreading the disease (<xref ref-type="bibr" rid="ref48">Ghorbani et al., 2024a</xref>). The susceptible host, the chili plant, and the virulent pathogen, <italic>Pythium</italic> spp., along with conducive environmental conditions, are key elements for disease establishment. Temperature, moisture, and organic matter (<xref ref-type="bibr" rid="ref54">Hansen and Keinath, 2013</xref>) have been directly linked with the incidence and severity of the damping-off owing to better establishment of the <italic>Pythium</italic> spp. with respect to attachment, establishment, and penetration into host tissues. Other environmental factors such as soil type, clay soil with high moisture holding capacity, rainfall intensity and duration, irrigation, humidity, and seedling surface witness are key factors responsible for damping-off disease. The pre-emergence damping-off requires 12&#x00B0;C while post-emergence damping-off require18-30&#x00B0;C and the optimum range lies from 24 to 30&#x00B0;C for successful infection of <italic>Pythium</italic> spp. (<xref ref-type="bibr" rid="ref31">Dubey et al., 2019</xref>). Furthermore, a high soil moisture (<xref ref-type="bibr" rid="ref104">Martin and Loper, 1999</xref>; <xref ref-type="bibr" rid="ref96">Majeed et al., 2019</xref>) supports the dissemination of propagules and increases the size of the spermosphere, while a pH of &#x02C3;5.8 supports successful infection. The relationship between environmental factors with inoculum and crop geometry spread also leads to the possible development of disease (<xref ref-type="bibr" rid="ref9">Arora et al., 2021</xref>). The detection and diagnosis of <italic>Pythium</italic> spp. pathogenic to chili plants involves utilizing both morphological characteristics and the analysis of specific DNA marker regions from the isolated pathogen. Recent research on <italic>Pythium</italic> spp. has predominantly focused on advanced molecular techniques based on PCR, including real-time PCR, multiplex PCR, loop-mediated isothermal amplification (LAMP) (<xref ref-type="bibr" rid="ref155">Sridapan and Krajaejun, 2023</xref>), and internal transcribed spacer (ITS) analysis (<xref ref-type="bibr" rid="ref43">Gaikwad et al., 2023</xref>).</p>
</sec>
<sec id="sec10">
<label>6.3</label>
<title>Infection stages and disease cycle</title>
<p><italic>Pythium</italic> spp. are eukaryotic organisms composed of filamentous, non-septate hyphae. Cell walls are primarily composed of cellulose and glucan, with limited amounts of chitin (<xref ref-type="bibr" rid="ref76">Kiselev, 2020</xref>). <italic>Pythium</italic> spp. in soil is attracted to the exudates released from germinating seeds and developing seedlings, leading to seed rot and seedling death (<xref ref-type="bibr" rid="ref177">Wohor et al., 2022</xref>). There are two main stages of <italic>Pythium</italic> spp. When the environmental conditions are favorable in the first stage, <italic>pyrium</italic> produces sporangia that grow into hyphae and vesicles with motile zoospores asexually. They are attracted to plant roots or other organic matter in the soil. In the other stage, <italic>Pythium</italic> reproduces sexually by combining an antheridium and an oogonium to form oospores, mostly during unfavorable environments. Sporangia can be single-celled or multi-celled and can be motile or non-motile.</p>
<p>Furthermore, <italic>Pythium</italic> spp. can be heterothallic or homothallic depending on the type of reproduction (<xref ref-type="fig" rid="fig1">Figure 1</xref>). When spores come into contact with a susceptible host, they can adsorb onto the root surface, forming a germ tube. The germ tube then penetrates the host&#x2019;s cells via wounds, root tips, or direct penetration, grows to branch mycelium through intracellular and intercellular spaces, and absorbs nutrients while exhibiting pathogenic symptoms externally (<xref ref-type="bibr" rid="ref81">Kushwaha, 2020</xref>). Sexual spores can thrive dormant in the soil in harsh environmental conditions as saprophytes and become active again when conditions become favorable (<xref ref-type="bibr" rid="ref126">Parveen et al., 2020</xref>).</p>
<fig position="float" id="fig1">
<label>Figure 1</label>
<caption>
<p>Infection stages and disease cycle of damping off disease.</p>
</caption>
<graphic xlink:href="fmicb-15-1479957-g001.tif"/>
</fig>
</sec>
</sec>
<sec id="sec11">
<label>7</label>
<title>Integrated management of damping-off</title>
<p>Chemical control methods, such as applying pesticides and other synthetic compounds, have been widely used for managing various pests and diseases. While initially effective, these methods often face significant challenges, including developing resistance in target species, environmental contamination, and negative impacts on non-target organisms, including beneficial species (<xref ref-type="bibr" rid="ref134">Ranjbar et al., 2023</xref>; <xref ref-type="bibr" rid="ref50">Ghorbani et al., 2024b</xref>). Over time, repeated use of chemical agents can lead to diminishing returns, requiring higher doses for the same effect and exacerbating ecological damage (<xref ref-type="bibr" rid="ref49">Ghorbani et al., 2023</xref>; <xref ref-type="bibr" rid="ref120">Nanehkaran et al., 2024</xref>). Additionally, concerns regarding human health risks and ecosystem disruption further highlight chemical control&#x2019;s limitations. These factors underscore the necessity of an integrated approach, combining biological, physical, and chemical methods to achieve more sustainable and resilient pest management strategies. Seed treatments are conducted to improve germination and seedling vigor (<xref ref-type="bibr" rid="ref19">Bhattarai et al., 2022</xref>). There are physical, chemical, and biological seed treatment methods in use aiming at seed disinfection, disinfestation, and seed protection. Chemical seed treatment includes soaking seeds in thiram, captan, and brassicol at the rate of 2.5&#x2009;g/kg (<xref ref-type="bibr" rid="ref111">Misra et al., 2022</xref>), while physical methods include hot air, hot water, and electron treatments, compressed moist heat (<xref ref-type="bibr" rid="ref140">Saha and Kumari, 2023</xref>). Biological seed treatment methods are neem leaf extract, garlic clove extract, ginger extract, and allamonda extract for chili (<xref ref-type="bibr" rid="ref110">Mishra et al., 2020</xref>). The promising microbial seed treatment is carried out by treating chili seeds with <italic>T. viride</italic> and <italic>P. fluorescence</italic> (<xref ref-type="bibr" rid="ref112">Mondal et al., 2022</xref>). Using synthetic seed treatments leads to phytotoxicity and germination defects, ultimately affecting human and animal safety. This is a disease preventive measure that can be taken before infestation from soil (<xref ref-type="bibr" rid="ref171">Volynchikova and Kim, 2022</xref>).</p>
<p>Good agricultural practices follow guidelines such as sterilizing nursery soil using a combination of physical and biological methods, including solarization and the application of <italic>Trichoderma</italic> as a soil drench (<xref ref-type="bibr" rid="ref80">Kumhar et al., 2022</xref>). Additionally, measures like burning soil with rice husks and bran, selecting soil from areas with no prior <italic>Pythium</italic> infestation history, using pure water and clean implements, using clean polythene sheets and organic amendments as mulch, fertilizing with organic materials from trustworthy, uncontaminated sources, and preventing water runoff from external sources are vital to avoid soil pathogen contamination (<xref ref-type="bibr" rid="ref184">Zamoum et al., 2022</xref>). If symptoms of damping-off arise, the immediate uprooting and isolation of affected seedlings from the rest of the plants is necessary to prevent further spread. These practices collectively contribute to the successful adoption of best cropping practices.</p>
<p>In situations where the aforementioned preventive measures prove insufficient due to the severity of the infestation, the use of chemical control becomes a viable option. It is crucial to first assess the extremity of the disease within the nursery to determine the appropriate dosage of fungicide required. Before moving to chemical intervention, it is imperative to consider the soil&#x2019;s overall health and the purity of the seeds selected for cultivation. More importantly, an immediate foundation of research outcomes to elucidate the morphology and physiology of <italic>Pythium</italic> and its response to external factors is needed to draw decisions. Fungicides should only be employed when all other methods have been exhausted, and the disease cannot be effectively managed or eradicated through alternative measures.</p>
<sec id="sec12">
<label>7.1</label>
<title>Use of chemical fungicides</title>
<p>Chemical control via fungicides is often considered a quick and effective way to prevent pathogens from infecting plant systems, particularly when applied to young, growing tissues like leaves, fruits, and flowers. This method is widely adopted due to the immediate results it yields compared to the lengthy process of developing resistant cultivars (<xref ref-type="bibr" rid="ref147">Sawant et al., 2022</xref>). Farmers have resorted to various chemical alternatives to methyl bromide for soil fumigation, as non-chemical methods can sometimes be labor-intensive and less effective against soilborne diseases (<xref ref-type="bibr" rid="ref32">Duniway, 2002</xref>). However, the use of chemical fungicides is constrained by a limited number of products registered for use on various crops and the high costs associated with these products, limiting chemical management of damping-off to a few active ingredients (<xref ref-type="bibr" rid="ref46">Garz&#x00F3;n et al., 2011</xref>). Fungicides from groups such as captan, benzimidazole, triazole, and dicarboximide have been recognized for their efficacy in controlling damping-off diseases (<xref ref-type="bibr" rid="ref82">Lamichhane et al., 2016</xref>). Specifically, metalaxyl, tridiazole, and captan are effective against <italic>Pythium</italic> and <italic>Phytophthora</italic> species, while maneb and mancozeb work against <italic>Pythium</italic> and <italic>Fusarium</italic> species, and these are commonly used by growers (<xref ref-type="bibr" rid="ref82">Lamichhane et al., 2016</xref>). Using systemic fungicides like metalaxyl before sowing can reduce <italic>Phytophthora</italic> and <italic>Pythium</italic> populations in the soil (<xref ref-type="bibr" rid="ref138">Russell et al., 1990</xref>). For seed treatments, Satija and Hooda found that benlate (0.1%) and dithane M-45 were the most effective against <italic>Pythium aphanidermatum</italic> and <italic>Fusarium solani</italic> (<xref ref-type="bibr" rid="ref6">Ansari et al., 1990</xref>). Soil drench applications of captan (<xref ref-type="bibr" rid="ref74">Khrieba, 2020</xref>), as well as soil treatments with methyl bromide, solarization, and metalaxyl soil drench, have been successful in preventing <italic>P. aphanidermatum</italic> infection (<xref ref-type="bibr" rid="ref57">Hickman and Michailides, 1998</xref>).</p>
</sec>
<sec id="sec13">
<label>7.2</label>
<title>Recommended cultural practices</title>
<p>Cultural practices are vital in ensuring disease-free chili production with optimum yield. In detail, proper site selection, avoiding areas with a history of damping-off infection, ensuring good drainage, applying organic amendments to enhance soil quality, adopting the best cropping systems, and using healthy seeds are best cultural practices to avoid damping-off disease (<xref ref-type="bibr" rid="ref93">Magnee et al., 2022</xref>). Further measures include infected plant parts being removed as soon as possible from the fields (<xref ref-type="bibr" rid="ref100">Manjunatha et al., 2022</xref>), timely and adequate fertilizing, irrigation, mulching to regulate soil temperature and suppress weeds, implementation strategies to protect crops from pests and diseases, sterilization through solarization or seed treatment (<xref ref-type="bibr" rid="ref8">Arora et al., 2022</xref>), cover cropping, green manuring, crop rotation, tillage, and managing proper space between crops (<xref ref-type="bibr" rid="ref113">Moura et al., 2022</xref>). Soil physiochemical properties management, such as pH, CEC, and moisture, is also important. Combining these practices is the most effective approach to managing damping-off disease in chili crops (<xref ref-type="bibr" rid="ref27">Cook and Haglund, 1991</xref>; <xref ref-type="bibr" rid="ref26">Cook, 2001</xref>). Furthermore, enhancing seed vigor is crucial for controlling damping-off, even in high pathogen density (<xref ref-type="bibr" rid="ref83">Lamichhane et al., 2017</xref>).</p>
</sec>
<sec id="sec14">
<label>7.3</label>
<title>Use of botanicals and biological control agents</title>
<p>The utilization of botanical extracts presents a highly promising approach for managing chili damping-off disease caused by <italic>Pythium</italic> spp. Plant extracts, including neem, garlic, ginger, turmeric, lemon, and pepper, have been shown to possess inherent antifungal properties. Consequently, these botanical extracts can be a viable and eco-friendly substitute (<xref ref-type="bibr" rid="ref129">Postma et al., 2003</xref>) for conventional chemical fungicides (<xref ref-type="bibr" rid="ref63">Islam and Faruq, 2012</xref>; <xref ref-type="bibr" rid="ref61">Hyder et al., 2021</xref>; <xref ref-type="bibr" rid="ref8">Arora et al., 2022</xref>). However, it is essential to recognize that the effectiveness of botanicals can show variability dependent upon the particular plant species, the specific plant part used, and the method that was used for extraction (<xref ref-type="bibr" rid="ref8">Arora et al., 2022</xref>). Moreover, the utilization of plant growth-promoting rhizobacteria (PGPR) in disease management endeavors shows promising results in the biocontrol of <italic>Pythium</italic> spp., as well as their positive impact on promoting the growth of solanaceous crops (<xref ref-type="bibr" rid="ref72">Kenawy et al., 2019</xref>), and many other major crop varieties. In many cases, it has been properly noted that imported bioformulations occasionally exhibit suboptimal performance owing to the influence of climate change (<xref ref-type="bibr" rid="ref25">Compant et al., 2010</xref>) and limitations in nutrient availability (<xref ref-type="bibr" rid="ref70">Kandeler et al., 2006</xref>). Therefore, the detection and characterization of PGPR endemic to chili rhizospheres capable of suppressing the inoculum of <italic>P. myriotylum</italic> and enhancing the growth of chilies in nurseries is in need.</p>
</sec>
<sec id="sec15">
<label>7.4</label>
<title><italic>Trichoderma</italic> and <italic>Bacillus</italic> as a biocontrol agent against <italic>Pythium</italic> spp</title>
<p><italic>Trichoderma</italic>, a prominent genus of filamentous fungi within the phylum Ascomycota, demonstrates remarkable potential as a biocontrol agent (BCA) against <italic>Pythium</italic> pathogens (<xref ref-type="bibr" rid="ref163">Thambugala et al., 2020</xref>; <xref ref-type="bibr" rid="ref180">Yao et al., 2023</xref>). These beneficial fungi, particularly the avirulent strains, have proven effective in plant protection, biostimulation, and biofertilization. Their effectiveness in agricultural applications depends on their metabolic activity and interactions with plants and other microorganisms.</p>
<p><italic>Trichoderma</italic> species can colonize the rhizoplane and plant roots (<xref ref-type="bibr" rid="ref130">Poveda and Eugui, 2022</xref>). They produce an array of metabolites with antimicrobial properties, including cell wall-degrading enzymes, both volatile and non-volatile antibiotics (<xref ref-type="bibr" rid="ref65">Jayaraj et al., 2006</xref>), as well as phytohormones and phytoregulators such as Indole Acetic Acid (IAA), cytokinin and ethylene that stimulate plant growth indirectly (<xref ref-type="bibr" rid="ref132">Rahman et al., 2012</xref>). They also secrete chitinase, which enhances plant defense mechanisms (<xref ref-type="bibr" rid="ref158">Subash et al., 2013</xref>) against fungal pathogens, and 1&#x2013;3 glucanase, a protease contributing to fungal pathogen defense and improved nutrient supply (<xref ref-type="bibr" rid="ref101">Mannai and Boughalleb-M&#x2019;Hamdi, 2023</xref>). Conversely, <italic>Trichoderma</italic> stimulates plant resistance locally or systemically by releasing products known as elicitors (<xref ref-type="bibr" rid="ref96">Majeed et al., 2019</xref>). These elicitors originate from the cell walls of both the host plant (endoelicitors) and the invading microorganism (exoelicitors). <italic>Pythium</italic> acidifies soil by secreting organic acids that can solubilize phosphate and other micronutrients supporting plant nutrition (<xref ref-type="bibr" rid="ref132">Rahman et al., 2012</xref>).</p>
<p>Notably, <italic>Trichoderma</italic> can control <italic>Pythium</italic> spp. through mechanisms such as necrotrophic mycoparasitism, competition for nutrients and space, and the synthesis of antifungal metabolites (<xref ref-type="bibr" rid="ref166">Ty&#x015B;kiewicz et al., 2022</xref>). These fungi exhibit rapid growth, metabolic versatility, and resistance to various toxic chemicals, including fungicides and herbicides, as adaptations (<xref ref-type="bibr" rid="ref33">El Enshasy et al., 2020</xref>; <xref ref-type="bibr" rid="ref66">Jia et al., 2024</xref>; <xref ref-type="bibr" rid="ref179">Yang et al., 2024</xref>).</p>
<p>Among <italic>Trichoderma</italic> species, <italic>T. harzianum</italic> strains stand out for their ability to protect plants by antagonizing <italic>Pythium</italic> pathogens in damping-off (<xref ref-type="bibr" rid="ref119">Muthukumar et al., 2011b</xref>), showing more promising performance than <italic>T. viridae, B. subtilis,</italic> and <italic>T. asperellum,</italic> which exhibit synergistic effects against <italic>P. aphanidermatum</italic> in solanaceous crops (<xref ref-type="bibr" rid="ref75">Kipngeno et al., 2015</xref>). Peroxidase activity was significantly higher in the roots of sugar beet seedlings treated with <italic>T. virens</italic> as biocontrol agents (<xref ref-type="bibr" rid="ref55">Hanson and Howell, 2004</xref>).</p>
<p><italic>Pseudomonas fluorescens</italic> is a beneficial bacterium widely used as a biocontrol agent against damping-off disease in chili via antibiosis, siderophore production, competition, induced systemic resistance, and enzyme production against damping-off disease in chili (<xref ref-type="bibr" rid="ref108">Mehmood et al., 2023</xref>). Bacterial species such as <italic>Bacillus</italic> have been proven to control fungal diseases. <italic>B. subtilis</italic> showed high antagonistic activity against <italic>Colletotrichum gloeosporioides</italic>, which caused anthracnose disease of chili (<xref ref-type="bibr" rid="ref10">Ashwini and Srividya, 2014</xref>). <italic>Bacillus</italic> has chitinolytic activity to control different fungal pathogens. Chitinolytic is a pathogenesis-related protein that increases the plant&#x2019;s defense mechanism against fungal pathogens (<xref ref-type="bibr" rid="ref91">Mabuchi et al., 2000</xref>; <xref ref-type="bibr" rid="ref58">Huang et al., 2005</xref>). <italic>Bacillus</italic> species were found to colonize the root surface, increase plant growth, and cause the lysis of fungal mycelia (<xref ref-type="bibr" rid="ref34">El-Bendary et al., 2016</xref>). As shown in previous reports, <italic>Bacillus</italic> spp. significantly control the damping-off chili caused by <italic>P. myriotylum</italic> (<xref ref-type="bibr" rid="ref67">Jimtha et al., 2016</xref>). Similar results from <xref ref-type="bibr" rid="ref61">Hyder et al. (2021)</xref> showed that <italic>B. cereus</italic> and <italic>B. megaterium</italic> significantly control damping off disease (<xref ref-type="bibr" rid="ref61">Hyder et al., 2021</xref>).</p>
</sec>
<sec id="sec16">
<label>7.5</label>
<title>Use of resistant varieties</title>
<p>Recent advances in plant breeding have significantly improved efforts to develop resistant chili cultivars to combat biotic and abiotic stresses. Marker-assisted selection (MAS) has accelerated the identification and incorporation of resistance genes into new cultivars, allowing breeders to develop varieties resistant to diseases such as powdery mildew, anthracnose, bacterial wilt, and viral diseases. Quantitative trait loci (QTL) mapping has been crucial in identifying resistance traits linked to these pathogens. Additionally, genomic selection and CRISPR-Cas9 gene editing technologies are revolutionizing breeding by enabling the precise modification of specific genes responsible for resistance. Traditional methods like germplasm screening and hybridization remain integral, but these are now supplemented with advanced genomic tools to improve efficiency and precision.</p>
<p>Host plants have evolved complex defense mechanisms that can either activate or suppress a range of genes when under attack by pathogens (<xref ref-type="bibr" rid="ref79">Kumar et al., 2022</xref>). These induced resistance responses include various physiological processes such as the generation of reactive oxygen species, synthesis of phytohormones, and production of pathogenesis-related (PR) proteins. The utilization of host plant resistance as a management strategy encompasses two distinct approaches: (i) the utilization of plant varieties that possess resistance to pathogens, resulting in reduced pathogen populations or enhanced ability to resist pathogen-induced damage, and (ii) the integration of these resistant varieties with other management tactics as part of an integrated pest management (IPM) framework. However, a significant number of plant diseases, especially those responsible for damping-off diseases, lack any plant cultivar that exhibits quantifiable resistance (<xref ref-type="bibr" rid="ref13">Babadoost and Islam, 2003</xref>). As a result, the optimal utilization of crop varieties possessing pathogen resistance can only be achieved by effectively integrating them with other strategies for disease management. However, there has been a lack of emphasis on incorporating plant resistance into other IPM strategies and a lack of research on quantifying the benefits of plant resistance within IPM programs that utilize various approaches (<xref ref-type="bibr" rid="ref29">Davis, 2009</xref>).</p>
<p>The current breeding methodology employed thus far for the development of crop varieties that exhibit resistance should be reevaluated if the intent is to prioritize sustainable crop protection strategies based on IPM. This is especially valid because the majority of cultivated plant varieties developed thus far have been rooted in a market-oriented strategy emphasizing the cultivation of high-yielding and economically advantageous crop varieties. The current tendency has resulted in increased utilization of short rotations or monoculture methods while neglecting the potential benefits that minor crops may offer for IPM (<xref ref-type="bibr" rid="ref109">Mess&#x00E9;an et al., 2016</xref>). Crop diversification is hindered by the limited range of minor crop types, which limits beneficial activities like multiple cropping and intercropping (<xref ref-type="bibr" rid="ref18">Benvenuto et al., 2020</xref>). Hence, in light of the pivotal role of breeding in enhancing crop competitiveness and facilitating adaptation to diverse cropping systems, it is imperative to use a distinct approach for breeding in IPM compared to conventional methods (<xref ref-type="bibr" rid="ref18">Benvenuto et al., 2020</xref>). Detailed information about management strategies is described in <xref ref-type="table" rid="tab4">Table 4</xref>.</p>
<table-wrap position="float" id="tab4">
<label>Table 4</label>
<caption>
<p>Control measures for managing damping-off in chili reported from different parts of the world.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="top">Species targeted</th>
<th align="left" valign="top">Active ingredients</th>
<th align="left" valign="top">References</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="top" colspan="3">Chemical control</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>Pythium aphanidermatum</italic>
</td>
<td align="left" valign="top">Propamocarb</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref97">Male and Vawdrey (2010)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. ultimumin</italic>
</td>
<td align="left" valign="top">Carbendazim and copper oxychloride</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref154">Singh et al. (2019)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. ultimumin</italic>
</td>
<td align="left" valign="top">metalaxyl/mefenoxam</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref20">Buchenau et al. (1981)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top"><italic>Pythium</italic> spp.</td>
<td align="left" valign="top">Metalaxyl, cyamoxanil, and carbendazim</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref182">Zagade et al. (2016)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. ultimum</italic>
</td>
<td align="left" valign="top">Metalaxyl</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref183">Zagade et al. (2014)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. ultimum</italic>
</td>
<td align="left" valign="top">Benlate</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref6">Ansari et al. (1990)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. ultimum</italic>
</td>
<td align="left" valign="top">Metalaxyl and Captan</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref47">Gholve et al. (2014)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. ultimum</italic>
</td>
<td align="left" valign="top">Carbendazim and Mancozeb</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref47">Gholve et al. (2014)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. aphanidermatum</italic>
</td>
<td align="left" valign="top">Thiram 75 WS and Captan 50 WP</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref141">Saha et al. (2011)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. aphanidermatum</italic>
</td>
<td align="left" valign="top">Metalaxyl</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref182">Zagade et al. (2016)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. ultimum</italic>
</td>
<td align="left" valign="top">Metalaxyl</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref182">Zagade et al. (2016)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. ultimum</italic>
</td>
<td align="left" valign="top">Ethaboxam</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref176">White et al. (2019)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. ultimum</italic>
</td>
<td align="left" valign="top">Ethaboxam</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref173">Wang et al. (2021)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. graminicola</italic>
</td>
<td align="left" valign="top">Carboxin and Thiram</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref31">Dubey et al. (2019)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top" colspan="3">Biological control</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. aphanidermatum</italic>
</td>
<td align="left" valign="top"><italic>T. viride</italic> and <italic>P. fluorescens</italic></td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref117">Muthukumar et al. (2011a)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. aphanidermatum</italic>
</td>
<td align="left" valign="top"><italic>T. viride</italic> and <italic>P. fluorescens</italic></td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref116">Muthukumar et al. (2010)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. aphanidermatum</italic>
</td>
<td align="left" valign="top">
<italic>Calothrix elenkenii</italic>
</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref99">Manjunath et al. (2010)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. ultimum</italic>
</td>
<td align="left" valign="top">
<italic>Stenotrophomonas rhizophila</italic>
</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref85">Lara-Capistran et al. (2020)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. ultimum</italic>
</td>
<td align="left" valign="top">
<italic>B. subtilis</italic>
</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref85">Lara-Capistran et al. (2020)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. debaryanum</italic>
</td>
<td align="left" valign="top">
<italic>B. subtilis</italic>
</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref5">Ali et al. (2016)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. aphanidermatum</italic>
</td>
<td align="left" valign="top">
<italic>T. harzianum</italic>
</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref107">Mehetre and Kale (2011)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. ultimum</italic>
</td>
<td align="left" valign="top">
<italic>Cryptococcus laurentii</italic>
</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref119">Muthukumar et al. (2011b)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. aphanidermatum</italic>
</td>
<td align="left" valign="top">
<italic>Streptomyces griseoviridis</italic>
</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref131">Punja and Yip (2003)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. ultimum</italic>
</td>
<td align="left" valign="top">
<italic>Gliocladium catenulatum</italic>
</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref106">Mcquilken et al. (2001)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. myriotylum</italic>
</td>
<td align="left" valign="top">
<italic>P. putida</italic>
</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref61">Hyder et al. (2021)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. aphanidermatum</italic>
</td>
<td align="left" valign="top">
<italic>Exiguobacterium indicum</italic>
</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref4">Al-Hussini et al. (2019)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top">
<italic>P. aphanidermatum</italic>
</td>
<td align="left" valign="top">
<italic>Actinoplanes campanulatus</italic>
</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref36">El-Tarabily et al. (2009)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top" colspan="3">Physical control</td>
</tr>
<tr>
<td align="left" valign="top"><italic>Pythium</italic> spp.</td>
<td align="left" valign="top">Tillage</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref15">Bailey and Lazarovits (2003)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top"><italic>Pythium</italic> spp.</td>
<td align="left" valign="top">Cover crops and soil residue management</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref15">Bailey and Lazarovits (2003)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top"><italic>Pythium</italic> spp.</td>
<td align="left" valign="top">Crop rotation and intercropping</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref59">Hwang et al. (2008)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top"><italic>Pythium</italic> spp.</td>
<td align="left" valign="top">Moderate humidity and avoid waterlogging, adequate light, and optimal temperatures</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref150">Schmidt et al. (2004)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top"><italic>Pythium</italic> spp.</td>
<td align="left" valign="top">Phosphorus, potassium, and calcium</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref84">Landis (2013)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top"><italic>Pythium</italic> spp.</td>
<td align="left" valign="top">Avoid excessive plant densities</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref84">Landis (2013)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top"><italic>Pythium</italic> spp.</td>
<td align="left" valign="top">Proper sowing date and good drainage</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref60">Hwang et al. (2000)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top"><italic>Pythium</italic> spp.</td>
<td align="left" valign="top">Soil pH</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref83">Lamichhane et al. (2017)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top"><italic>Pythium</italic> spp.</td>
<td align="left" valign="top">Seedbed preparation</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref124">Njoroge et al. (2008)</xref>
</td>
</tr>
<tr>
<td align="left" valign="top"><italic>Pythium</italic> spp.</td>
<td align="left" valign="top">Seed quality</td>
<td align="left" valign="top">
<xref ref-type="bibr" rid="ref136">Roberts et al. (2016)</xref>
</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="sec17">
<label>7.6</label>
<title>Detrimental effects of synthetic fungicides</title>
<p>Even though chemical fungicides are effective in rapidly controlling damping-off, they lead to human and ecotoxicity, particularly in developing nations (<xref ref-type="bibr" rid="ref172">Voorrips et al., 2004</xref>). Human toxicity includes irritant dermal injuries, mucus membranes, and dermal sensitization. For animals, livestock poisoning and aquatic organisms, particularly fish, experience adverse impacts from fungicides. Furthermore, fungicides reduce the fungal and total microbial biomass in soil (<xref ref-type="bibr" rid="ref170">Verdenelli et al., 2023</xref>), influencing the imbalance in soil microbiota, and another detrimental effect is phytotoxicity by fungicide residues (<xref ref-type="bibr" rid="ref77">Kova&#x010D;i&#x010D; et al., 2013</xref>). Different fungicides have different modes of action and differ in how long they last to suppress disease. At the same time, fungicide resistance occurs with the longer usage of the same fungicide type. For a greater chance of enhanced disease protection in the fields, it is strongly advised to rotate two or more distinct classes of fungicides to prevent the development of fungicide-resistant pathogens (<xref ref-type="bibr" rid="ref41">F&#x00F6;rster et al., 2007</xref>). It is hard to achieve sustainable disease management by using fungicides alone. Hence, the need for integrated disease management is discussed.</p>
</sec>
<sec id="sec18">
<label>7.7</label>
<title>Advances in molecular aspects of detection as well as management of damping-off diseases</title>
<p>Recent advances in the molecular understanding of damping-off diseases have significantly enhanced detection and management strategies. Molecular diagnostics, such as polymerase chain reaction (PCR) and quantitative PCR (qPCR), now allow for rapid and precise identification of soil-borne pathogens like <italic>Pythium</italic>, <italic>Fusarium</italic>, and <italic>Rhizoctonia</italic> at early stages of infection. These techniques enable species-level identification and quantification of pathogen load in soil and plant tissues, which can help predict disease outbreaks. The advent of metagenomics and next-generation sequencing (NGS) has further revolutionized detection by allowing a comprehensive analysis of microbial communities, identifying emerging pathogens, and understanding the microbial shifts that occur during disease progression (<xref ref-type="bibr" rid="ref42">Fu et al., 2024</xref>).</p>
<p>On the management side, molecular advances have improved the use of biological control agents and the development of resistant cultivars. Functional genomics, through tools like RNA interference (RNAi) and CRISPR-Cas9, is used to knock out or modify plant susceptibility genes, providing a novel method for increasing disease resistance. Furthermore, insights into the molecular interactions between pathogens and biocontrol agents, such as <italic>Trichoderma</italic> and <italic>Pseudomonas fluorescens</italic>, have led to enhanced formulations that can better suppress pathogen activity through targeted modes of action, such as mycoparasitism and antibiosis. These advances highlight the critical role of molecular tools in both early detection and integrated management approaches, offering a promising future for sustainable control of damping-off diseases.</p>
</sec>
</sec>
<sec sec-type="conclusions" id="sec19">
<label>8</label>
<title>Conclusion</title>
<p>Damping-off disease remains a formidable challenge in chili cultivation, further exacerbated by environmental changes and the advent of more aggressive pathogen strains. Our comprehensive examination reveals that integrated disease management is the key to addressing this issue. While chemical fungicides provide immediate control, their long-term usage is unsustainable due to environmental and human health concerns as well as the potential for resistant pathogen strains to develop. Sustainable management of damping-off requires a multifaceted approach, including the development of resistant chili varieties, the implementation of strategic cultural practices, and the adoption of novel technologies such as nanoparticles. These strategies must be adaptable to evolving climatic conditions and tailored to local agro-ecological contexts.</p>
<p>Moreover, there is a pressing need for further molecular studies to enhance our understanding of plant&#x2013;microbe interactions, which could lead to more effective and targeted control measures. The innovative application of nanoparticles in the laboratory setting shows great promise and could pave the way for new biocontrol methods that are both effective and environmentally sustainable. Ultimately, a comprehensive integrated pest management framework that combines traditional practices with cutting-edge research is essential for the holistic and sustainable management of damping-off disease in chili cultivation. As we advance our scientific knowledge and integrate it into field-based applications, we move closer to safeguarding the future of chili production against the persistent challenges posed by this destructive disease.</p>
<sec id="sec20">
<label>8.1</label>
<title>Future prospects</title>
<p>The challenge of damping off disease remains as critical as ever, especially with its large-scale spread exacerbated by flooding and climate change. Despite extensive research on the epidemic nature of the disease, gaps still persist in our understanding of host&#x2013;pathogen interactions, disease proliferation, and effective management strategies.</p>
<p>There is an urgent need to devise efficient integrated management strategies that consider the increasing incidence of flooding, environmental factors, and the variability of pathogenic races. Developing resistant varieties of chili appears to be one of the most promising approaches for the long-term management of damping-off disease. By breeding for resistance, we can reduce the vulnerability of crops to this disease under various environmental conditions.</p>
<p>In addition to developing resistant varieties, integrating cultural practices tailored to specific climatic changes is crucial for more effective disease management. As the climate continues to change, previously effective strategies may need to be adapted to remain effective under new conditions. Furthermore, there is a growing need for more molecular studies to deepen our understanding of plant&#x2013;microbe interactions. Insights into the infection mechanisms of pathogens will be invaluable for creating targeted management strategies that can interrupt the infection process and mitigate the disease&#x2019;s impact.</p>
<p>Additionally, our laboratory is exploring the potential of nanoparticles as a novel approach to combat <italic>Pythium</italic> spp. Nanoparticles represent an innovative frontier in managing damping-off disease in chili. These microscopic particles could offer new mechanisms of action against the pathogen, potentially enhancing the effectiveness of existing treatments or providing alternative management solutions. As this research advances, it could open new pathways for controlling this persistent and damaging disease.</p>
</sec>
</sec>
</body>
<back>
<sec sec-type="author-contributions" id="sec21">
<title>Author contributions</title>
<p>CD: Writing &#x2013; original draft. GM-U-D: Writing &#x2013; original draft. RA: Writing &#x2013; review &#x0026; editing, Data curation. TT: Writing &#x2013; review &#x0026; editing, Formal analysis, Data curation. RL: Writing &#x2013; review &#x0026; editing, Data curation, Conceptualization. YX: Writing &#x2013; review &#x0026; editing, Data curation, Conceptualization. SM: Writing &#x2013; review &#x0026; editing, Software, Conceptualization. AG: Writing &#x2013; review &#x0026; editing.</p>
</sec>
<sec sec-type="funding-information" id="sec22">
<title>Funding</title>
<p>The author(s) declare that financial support was received for the research, authorship, and/or publication of this article. We are thankful to the projects of the Longdong University Doctoral Fund Project (XYBYZK2108), Gansu Provincial Natural Science Foundation Project (22JR5RM207), and Gansu Provincial University Youth Doctoral Fund Project (2021QB-120).</p>
</sec>
<sec sec-type="COI-statement" id="sec23">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec sec-type="disclaimer" id="sec24">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<sec sec-type="supplementary-material" id="sec25">
<title>Supplementary material</title>
<p>The Supplementary material for this article can be found online at: <ext-link xlink:href="https://www.frontiersin.org/articles/10.3389/fmicb.2024.1479957/full#supplementary-material" ext-link-type="uri">https://www.frontiersin.org/articles/10.3389/fmicb.2024.1479957/full#supplementary-material</ext-link></p>
<supplementary-material id="SM1" xlink:href="Image_1.JPEG" mimetype="image/jpeg" xmlns:xlink="http://www.w3.org/1999/xlink">
<label>Supplementary Figure S1.</label>
<caption>
<p>The botanicals characteristics of chilli plant. <bold>(a)</bold> Chilli plant. <bold>(b)</bold> Ripe red fruit. <bold>(c)</bold> Immature green fruits. <bold>(d)</bold> Ripe red fruit.</p>
</caption>
</supplementary-material>
<supplementary-material id="SM2" xlink:href="Image_2.JPEG" mimetype="image/jpeg" xmlns:xlink="http://www.w3.org/1999/xlink">
<label>Supplementary Figure S2.</label>
<caption>
<p>Characteristics symptoms of damping-off disease on young seedlings of chilli plants. <bold>(a)</bold> Symptoms on young seedlings. <bold>(b)</bold> Whitish growth on the stem near soil surface. <bold>(c)</bold> Diseased plants wilted after severe attack. <bold>(d)</bold> Healthy plants.</p>
</caption>
</supplementary-material>
</sec>
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