Use of semiochemicals in integrated pest management

Introduction

Semiochemicals are chemical compounds or mixtures released by one organism that elicit behavioural or physiological responses in members of the same species or different species (Nordlund and Lewis, 1976) [73] .The adverse effect of pesticides on soil, water, humans, animals and ultimately the environment diverted the attention of the scientific community to develop an ecofriendly approach for managing the devastating crop pests (Manda et al., 2020) [60] . Semiochemicals can use in integrated pest management (IPM). These compounds play a crucial role in insect communication by mediating both insect-insect and plant-insect interactions. Chemical signals exchanged among insects, host plants and their environment are essential for insect survival enabling them to modify their behaviour in response to environmental cues. These cues known as semiochemicals influence physiological and behavioural responses within a species as well as between different species. Host plant volatiles serve as important semiochemical cues by helping insects locate essential resources including food, mates, oviposition sites and shelter (Prokopy et al., 1984; Witzgall et al., 2010) [80,112] . The term semiochemical originates from the Greek word "semeon" (Firn, 2011) [38] . Depending on the nature of the interactions, chemical communication between organisms can be broadly classified into two basic types: intraspecific and interspecific communication. Pheromones and allelochemicals are examples of intraspecific and interspecific chemical communication respectively (Nordlund et al., 1981) [74] . Pheromones have evolved as chemical signals that facilitate communication related to sexual attraction, alarm, aggregation, and territorial marking. In contrast, allelochemicals are classified based on who benefits from the interaction: kairomones benefit the receiver, allomones benefit the emitter, and synomones provide advantages to both organisms. The interactions between plant semiochemicals and insect pheromones differ in their modes of action, and their combined use can improve the effectiveness of pest management strategies (Bakthavatsalam and Subaharan, 2016) [11] . Furthermore, semiochemicals are categorized into several functional groups according to the type of information they convey and the ecological relationship between the emitting organism and the receiving organism during chemical communication (Vilela and Della Lucia, 2001) [109] . Semiochemicals play a crucial role in regulating insect behavior and are involved primarily in insect-insect and plant-insect interactions (Abd El Ghany, 2019) [1] . Because of their ability to modify pest behavior semiochemicals have become important tools in environmentally sustainable pest management programs (Howse et al., 1998) [45] . Semiochemicals are used by insects to mediate crucial behaviours like mating, oviposition and resource-seeking (Evenden and Silk, 2016) [37] .

A) Pheromone

Pheromones are chemical substances secreted by an organism that evoke a specific behavioural or physiological response in another individual of the same species (Karlson and Luscher (1959) [51] . They represent one of the most extensively studied classes of semiochemicals involved in intraspecific communication. The concept of chemical communication within the same species was initially described by Bethe (1932) [17] , who introduced the term "ectohormone". Later, Karlson and Butenandt (1959) [50] coined the term pheromone from the Greek "pherein" meaning "to transfer" and "hormon" meaning "to excite") to describe chemicals released by one individual and received by another individual of the same species producing a specific adaptive response. Pheromones play a vital role in regulating reproductive behaviour, aggregation, trail following, alarm signalling, territorial marking, caste differentiation and social organization in insects (Nordlund et al., 1981; Wyatt, 2014) [74,116] . In insects, pheromones are detected by specialized olfactory sensilla located mainly on the antennae and the resulting signals influence behaviours essential for survival and reproduction. A well-known example is the Queen Mandibular Pheromone (QMP) of the honey bee (Apis mellifera L.) whose major component is 9oxo-2-decenoic acid (9-ODA) suppresses ovarian development in worker bees, maintains colony organization and simultaneously functions as a sex attractant during the nuptial flights of virgin queens (Regnier, 1968; Winston, 1987) [87,111] . Similarly, sex pheromones released by female moths attract conspecific males over long distances, facilitating successful mating and species-specific communication (Carde and Baker, 1984; Wyatt, 2014) [116] .

Classes of Pheromones Based on the responses elicited pheromones can be classified into 2 groups-Releaser Pheromones: Rapid Behavioral Responses

Releaser pheromones are the most widely recognized class of pheromones because they trigger immediate and specific behavioral reactions in members of the same species (Karlson and Luscher, 1959) [51] . These responses are typically instinctive, stereotyped and mediated directly through the central nervous system allowing the chemical signal to rapidly initiate a particular behavior (Wyatt, 2003) [115] .

Primer Pheromones: Long-Term Physiological Effects

Unlike releaser pheromones, primer pheromones produce gradual changes in the recipient's physiological or endocrine system, resulting in delayed but sustained behavioral effects (Karlson and Luscher, 1959) [51] . Rather than provoking an immediate response, they regulate developmental and reproductive processes and play a vital role in maintaining social organization and coordinating the activities of members within a social group.

Pheromone reception: Insects are exceptionally skilled at chemical analysis, detecting and interpreting their surroundings through semiochemicals with remarkable sensitivity. For instance, a male moth can sense the sex pheromones released by a female even when they are present in extremely low concentrations. The antennae function as biological signal converters, transforming chemical cues into electrical nerve impulses that the brain can understand. These signals are detected by specialized sensory structures called sensilla, which are primarily located on the antennae and are then transmitted to the brain for further interpretation. Chemical ecologists take advantage of this highly sensitive olfactory system by using insect antennae as biological sensors to identify pheromones and other semiochemicals. Moreover, the insect olfactory system is highly selective, capable of distinguishing natural pheromones from compounds that differ only slightly in their chemical structure (Leal, 2005) [59] . Pheromones may be released by certain insects directly from their glands, while others do so use specialized structures like pheromone brushes or pouches (Mori, 1981) [69] .

Types of Pheromones 1) Sex pheromone

A sex pheromone is a semiochemical released by an organism (usually female) to attract an individual of the opposite sex (male) of the same species for mating or other activity related to reproduction (Landolt, 1997) [57] . Over 150 species of insects are known in which the females produce sex pheromones and about 50 species in which the males do so. It is useful in IPM for monitoring, mass trapping and mating disruption (Bakthavatsalam, 2016) [10] . Studies of several Hemiptera (true bugs) have mostly identified male-specific sex pheromones, which may represent historical pre judice rather than taxonomic class bias (Gomez-Diaz, 2013) [42] . Sex pheromones have been identified in more than 1600 insect species and are among the most widely studied semiochemicals because of their high species specificity and practical applications in integrated pest management (IPM). The use of synthetic sex pheromones provides an environmentally safe and speciesspecific alternative to conventional insecticides, thereby reducing pesticide use and preserving natural enemies (Witzgall et al., 2010; Baker, 2016; El-Sayed, 2022) [9,36,112] . A successful application of sex pheromones is the management of codling moth, Cydia pomonella Linn. in apple and pear orchards. Synthetic codlemone is released through pheromone dispensers to disrupt male orientation towards females, thereby reducing mating, oviposition and larval infestation. This mating-disruption technique has been successfully adopted as an important component of IPM and reduces dependence on conventional insecticides (Witzgall et al., 2008) [113] .

2) Aggregation pheromones

Aggregation pheromones attract individuals of both sexes to food sources, reproductive habitats, or shelter sites. Xiu et al. (2024) [117] reported that the aggregation pheromone of the stick tea thrips, Dendrothrips minowai Priesner (Thysanoptera: Thripidae), increased the capture of adults by 1.2-3.0 times on sticky traps compared with unbaited traps. Furthermore, sticky traps supplemented with the aggregation pheromone of D. minowai attracted significantly more adults, capturing 1.2-3.0 times as many individuals as control traps. In the Colorado potato beetle, the male-produced aggregation pheromone 'Decemdiol', is highly attractive to both sexes. In beetles, aggregation pheromones have also been identified whereas, in the predatory stink bug (Podisus spp.) they have not yet been identified, suggesting opportunities for developing push-pull systems for behavioural control (Jermy and Butt, 1991) [47] . Over recent decades, the importance of applying aggregation pheromones in the management of the boll weevil (Anthonomus grandis B.), stored product weevils (Sitophilus zeamais M., Sitophilus granaries L., Sitophilus oryzae L.) and pea and bean weevil (Sitona lineatus L.) has been demonstrated (Landolt, 1997) [57] . Ferrugineol-based aggregation pheromone traps used against red palm weevil attract both sexes and are used for mass trapping. Field demonstrations in Goa resulted in about 64 percent reduction in pest damage, demonstrating successful application of aggregation pheromones in IPM (Maruthadurai and Ramesh, 2020) [62] .

3) Alarm pheromones

Alarm pheromones exhibit considerable diversity among insects and function to warn conspecific individuals of potential danger or predator attack (Wyatt, 2003; Verheggen et al., 2010) [108,115] . In honey bees, alarm pheromones are produced by the sting apparatus and released during stinging thereby, alerting nearby colony members and triggering defensive behaviour. Similarly, ants utilize trail pheromones secreted from the Dufour's gland to mark pathways that guide nestmates to food sources and back to the nest, ensuring efficient communication and coordinated foraging within the colony (Bruce, T. J. A., 2015) [21] . Alarm pheromones may also function as kairomones by providing chemical cues that attract predators or parasitoids, thereby benefiting the receiver while adversely affecting the emitting organism (Nordlund and Lewis, 1976) [73] . Aphid alarm pheromones are among the most extensively studied examples. The alarm pheromone (E)-β-farnesene has been identified in several aphid species including Acyrthosiphon pisum H. (pea aphid), Aphis gossypii G. (cotton aphid), and Rhopalosiphum padi L. (bird cherry-oat aphid) (Bowers et al., 1972) [19] . Honey bees also release alarm pheromones from their stingers when defending the colony against organisms that threaten the hive (Boch et al., 1962; Shearer and Boch, 1965) [18,96] . Apthena (Anonymous, 2026) [7] is an innovative semiochemical-based product containing (E)-βfarnesene (EBF), an aphid alarm pheromone. It attracts beneficial predators and parasitoids, thereby enhancing natural biological control of aphids and supporting integrated pest management (IPM) while reducing dependence on synthetic insecticides.

4) Trail pheromone

Trail pheromones play a crucial role in the foraging behaviour of social insects, particularly ants (Morgan, 2009) [68] . By depositing trail pheromones, ants establish chemical pathways that guide other colony members to food sources and back to the nest. The characteristics of trail pheromones vary among species and are influenced by their specific habitats and ecological requirements (Attygalle and Morgan, 1985) [8,68] . Synthetic trail pheromones have been incorporated into baits and traps to selectively target pest ant species, demonstrating the strong and consistent influence of these chemical signals on ant behaviour in practical, largescale pest management applications (Vander Meer, 1996; Rust et al., 2004) [89,106] . Trail pheromones have been investigated for manipulating the foraging behaviour of termites. In Macrotermes annandalei S., the compound (Z)dodec-3-en-1-ol functions as a trail-following pheromone and induces orientation and recruitment behaviour. Synthetic pheromone trails were able to compete with natural trails under field conditions, demonstrating their potential for developing pheromone-based termite management strategies (Sillam-Dusses et al., 2001) [97]

5) Host marking pheromone

In several phytophagous insect species, females exhibit oviposition behavior by avoiding hosts that have already been marked with host-marking pheromones (HMPs). These pheromones function as chemical stimuli detected through chemoreception, enabling females to differentiate between previously marked and unmarked hosts. Such stimuli are classified as either signals or cues (Seeley, 1989) [92] . Chemical cues, for example, can indicate the presence of eggs or larvae already occupying a host (Butkewich et al., 1987) [3] . Field application of glutathione and glutamic acid, identified as HMPs of Ceratitis cosyra and C. rosa, respectively, reduced fruit-fly infestation in mango by approximately 70-75 percent compared with untreated controls. This demonstrates the potential of HMPs as a nontoxic, behaviour-based component of IPM (Cheseto et al., 2023) [25] .

B) Allelochemicals

Allelochemicals such as protease inhibitors, α-amylase inhibitors, lectins, tannins, phenolic compounds and flavonoids disrupt the digestive physiology of insects by affecting key gut enzymes. Protease inhibitors reduce protein digestion by inhibiting proteolytic enzymes whereas, α-amylase inhibitors prevent efficient starch digestion, resulting in nutrient deficiency and physiological stress in insects (Franco et al., 2002; Zavala et al., 2004) [40,121] . According to Norduland et al. (1981) allelochemicals can be separated into signals that are beneficial to the receiver (kairomones), the emitter (allomones) or both (synomones).

1) Allomone

Allomone was proposed by Brown and Eisner (1970) [20] to denote those substances which convey an advantage upon the emitter. Allomone is a chemical substance produced and released by one organism that affects the behavior or physiology of an individual of another species in a way that benefits the emitter, while the receiving organism gains no benefit and may even be harmed (Eisner and Meinwald, 1995) [35] . Different types of behaviours are mediated by allomones. Key among these are their roles as defensive secretions against predators or other natural enemies, such as the formic acid sprayed by formicine ants and the venom from many social wasps, social bees, and ants (Yusuf, 2021) [119] . For example, Defensive chemicals released by stink bugs or bombardier beetles that repel predators. Bark beetles communicate via pheromones to announce a new food resource (i.e. dead trees, roots, living trees, etc.) ultimately resulting in the accumulation of a large concentration of bark beetles. Select species of bark beetles have the capability to emit pheromones that can negatively affect the behavioral response of another competing species of bark beetles when both species are attempting to inhabit the loblolly pine tree (Wood, 1982) [114] .

2) Kairomone

Kairomones are interspecific semiochemicals or mixtures of semiochemicals produced by one species that trigger responses beneficial to an individual of another species receiving the signal. They are important in biological control programs because of the significant role of semiochemicals in host-plant selection by phytophagous insects (Bernays, 1994) [15] . Kairomone based lures are applied in crop systems, where the surrounding plant odours influence the ability of natural enemies to locate herbivorous insects (Schroder and Hilker, 2008; Randlkofer et al., 2010) [44, 84, 91] .

3) Apneumones

Chemical compounds released by non-living materials that trigger beneficial behavioural or physiological responses in one organism while potentially causing harmful effects to other species interacting with the same material. A rare example is the release of hexanal and 2-methyl-2-butanol from rabbit feces, which attracts female sandflies seeking suitable sites for egg-laying. These types of allelochemicals demonstrate the complexity of chemical communication among species, producing interactions that can range from mutually beneficial to detrimental depending on the organisms involved (Abd El-Ghany, 2020) [2] . Rare instances of these allelochemicals have been documented in the literature, such as the attraction of sandfly females for oviposition by hexanal and 2-methyl-2-butanol emitted by rabbit faeces (Dougherty et al., 1995) [32] .

1) Synomones

A synomone is an interspecific chemical signal that is beneficial to both interacting organisms, namely the emitter and the receiver. For example, the floral synomone of certain Bulbophyllum species (Orchidaceae) attracts male fruit flies (Tephritidae: Diptera), which act as pollinators. The orchid flowers are successfully pollinated, while the male Dacini fruit flies obtain a sex pheromone precursor or enhancer. The floral synomone also serves as a reward to pollinators and occurs in the form of phenylpropanoids, such as methyl eugenol (Tan et al., 2002; Nishida et al., 2004; Tan et al., 2006) [72] or phenyl butanoids, such as raspberry ketone (Tan and Nishida, 2000) and zingerone (Tan and Nishida, 2005). They are often released in response to herbivore attack, attracting natural enemies of herbivores and thereby reducing plant damage (Karban and Baldwin, 1997; Dicke and Hilker, 2003) [30,44,49] . These cues are utilized by parasitoids belonging to the families Eulophidae, Scelionidae, and Mymaridae in various ecosystems involving both arboreal perennial plants and herbaceous annual plants (Hilker and Meiners, 2002; Colazza et al., 2004) [26,44] . Plant-derived synomones released by maize following Chilo partellus S. damage, particularly (E)-β-farnesene and (Z)-3-hexenyl acetate, attract the parasitoid Cotesia flavipes C., thereby enhancing indirect plant defence and offering potential for parasitoidbased pest management (Ngumbi et al., 2005) [71] .

Use of Semiochemicals In Integrated Pest Management 1) Mass trapping

Mass trapping is an important component of integrated pest management (IPM) for managing soil-dwelling scarab beetles by suppressing adult populations and consequently reducing larval infestations. After successful preliminary experiments, large-scale mass trapping of the American palm weevil was initiated in 1991 on two major oil palm plantations in Costa Rica, encompassing areas of 6514 ha and 8719 ha, respectively (Oehlschlager et al., 2002) [75] . Galko et al., (2016) [41] evaluated the effectiveness of six pheromone-based traps for managing the European spruce bark beetle. The study compared four commercially available traps with two newly developed designs: BEKA (B), Ecotrap (E), Cross trap (K), Lindgren funnel trap (L), Funnel trap (P), and Theysohn (T). Differences in trapping performance were primarily attributed to variations in trap design. Field evaluations were carried out across ten locations over three consecutive growing seasons. The strengths and limitations of each trap type, identifying operating lifespan and storage durability as the major drawbacks affecting their practical use. Methyl eugenolbaited PAU fruit-fly traps used for managing Bactrocera spp. in guava during 2017 and 2018 in Punjab. The traps recorded peak catches of 801.5 and 720.4 flies per trap per week, respectively. Fruit infestation was reduced from 67-81 percent in untreated plots to 45-48 percent in trapped plots, while fruit yield increased from 8.1-8.8 to 12.6-15.4 kg per tree (Sharma et al., 2022) [95] . The Japanese beetle, Popillia japonica N. (Coleoptera: Scarabaeidae), has recently been effectively controlled using this approach (Pinero and Dudenhoeffer, 2018). Mass trap synthetic blend of 3Z-hexen-1-ol, α-pinene, hexyl acetate, benzyl alcohol, and 6Z nonenal in funnel traps yielded the highest capture of adult Mass trap synthetic blend of 3Zhexen-1-ol, α-pinene, hexyl acetate, benzyl alcohol, and 6Z nonenal in funnel traps yielded the highest capture of adult Diaphania indica Saunders (Lepidoptera: Crambidae) in snakegourds (Debnath et al., 2023) [28] . Pawar et al., (2009) [77] evaluated a sex pheromone-based IPM strategy against Leucinodes orbonalis in brinjal at MPKV, Rahuri. Mass trapping using Leucinlure-baited funnel traps at 1 trap per 100 square meter, integrated with cultural and botanical practices, reduced shoot infestation by 80.44 percent compared with the untreated control and 61.64 percent compared with IPM without mass trapping. The treatment also increased yield by 44.75 percent over the untreated control, demonstrating the effectiveness of pheromonebased mass trapping [82] Funnel Traps: These traps contain a pheromone lure and a funnel-shaped entrance that guides insects into a collection chamber, preventing their escape. They are suitable for capturing large numbers of insects (Raj, 2025) [82] .

Water Traps: Insects are attracted by the pheromone lure and fall into a container filled with water, where they drown (Raj, 2025) [82] .

Sticky Traps: These traps have an adhesive-coated surface that captures insects when they land on or come into contact with it. They are commonly used for pest monitoring (Raj, 2025) [82] .

Mass Trapping Devices: These are larger traps designed to capture a high number of insects, helping to reduce pest populations rather than only monitoring their presence (Raj, 2025) [82] 1) Mating disruption Mating disruption is a semiochemical-based pest management strategy that interferes with the pheromone communication system of insects, preventing successful mate location and copulation (shani, 2001) [93] Mating disruption was first discussed by the scientists of an institute in Bordeaux, France in 1974. These pheromones confuse individuals, hinder their ability to locate mates, and reduce successful mating, ultimately lowering reproductive potential (Carde and Minks, 1995). This technique relies on artificially synthesized pheromones, usually consisting of blends of major and minor chemical components, designed to imitate the activity of naturally produced pheromones (Bakthavatsalam, 2016) [10] . Various mating disruption mechanisms have been postulated (Bartell 1982; Carde and Minks 1995; Miller et al. 2006) [13,67] , which can be divided into two main categories: competitive and non-competitive (Miller et al., 2006) [67] . Competitive disruption includes competitive attraction (false-trail-following, confusion), induced allopatry, and induced arrestment, whereas noncompetitive disruption involves other mechanisms, such as suppressed calling/mating, camouflage, sensory imbalance, induced allochrony, and desensitisation (habituation) (Miller and Gut 2015) [66] . Mating disruption devices, several microencapsulated dispensers, manually applied dispensers (pheromone traps, PB rope, SPLAT), and high-emission dispensers are now available in the market. Consequently, this innovative technology may assist in addressing the gaps in current IPM programmes for sustainable pest management (Mevada et al., 2023) [63] . In overall, MD represents a key behavioral component in reducing pesticide reliance and promoting sustainable management of stored-product pests. (Adamic Zamljen, 2026) [5,120] . Mating disruption technology has been commercially adopted and is being increasingly utilized for the management of the vine mealybug, Planococcus ficus S., while its application against the California red scale, Aonidiella aurantii M., remains comparatively limited (Franco, 2022) [39] . The strategy is based on the use of electrostatically charged powders, able to adhere to insect cuticles, as carriers for active ingredients such as pheromones, entomopathogens, and insecticides (Trematerra, 2012) [105] . Paddy yellow stem borer Scirpophaga incertulas Walke(Lepidoptera: Crambidae) are effectivelymanaged by the Pheron RSB mating distruption dispenser (Iqbal et al., 2023) [46] . The mating disruption technique was used in pea fields, for five years period to control pea moth, Cydia nigricana F. by assessing male attraction to monitor traps, pod infestation, and larval age structure in pheromone-treated and untreated grain. Cellulose pheromone dispensers were manually attached to the top shoots of pea plants and released 540 mg per hectare per day synthetic pheromone (E8, E10-dodecadien-1-yl acetate). The dispensers had a half-life of about 30 days. Although male attraction to pheromone monitoring traps was largely suppressed at the edges and within MD fields (Saucke et al., 2014) [90] . PB-Rope L sex pheromone dispensers were used for mating disruption of pink bollworm in cotton. The treatment reduced male moth catches by 79.4 and 93.8 percent during two successive seasons and reduced boll damage and larval populations, while increasing seed-cotton yield by 32.1 and 40.9 percent, respectively (Patil et al. (2007) [76] .Yeast-derived sex pheromones used for mating disruption of Helicoverpa armigera in cotton. Three years of field trials in Greece showed that yeast-produced pheromones were as effective as synthetic pheromones in disrupting male orientation (Raptopoulos et al., 2025) [85] . Ricciardi et al. (2024) [4] evaluated a biodegradable double-tube sex pheromone dispenser (Isonet® L CG-BIOX235) for simultaneous mating disruption of Lobesia botrana and Cryptoblabes gnidiella in Italian vineyards. Field trials conducted in Tuscany and Apulia during 2022-2023 showed that the dispenser significantly reduced infestation of both pests compared with untreated vineyards.

2) Monitoring

One of the most common and successful practical applications of semiochemicals is in pest detection and monitoring (Witzgall et al., 2010) [112] . Pheromone traps can detect insect pests and are also valuable for estimating population density and seasonal fluctuations to identify the initial peak of flight activity (Weinzierl et al. 2005) [110] . Rhainds et al. (2016) [88] evaluated the abundance of male Choristoneura fumiferana C. (Lepidoptera: Tortricidae) in Canada using a surveillance method based on pheromonebaited traps. Variability in trap catches and its association with later infestations was utilized to develop a decisionsupport model (Bruce and Smart, 2009) [22] . New technologies have developed in the recent years for pest detection and surveillance systems, based on acoustic transducers that are capable to detect the sounds of the insects and then send electronic signals to a central station where the trap locations are identified (Beroza, 2002) [16] . Other systems based on an automatic counting of the captured insects fitted with crossing transducers integrated with data transmission systems have been created (Tabuchi et al., 2006) [98] . In an interesting different method, Guarnieri et al., (2011) [43] developed an automated electronic trap for monitoring the codling moth by altering a commercial trap-specific for the insect with the application of a mobile phone camera and transmit the data to a distant server. This type of technology is likely to be most financially feasible for detection of important exotic pests where the initial incursion of a new moth is the most valuable task. IoT-based traps with accurate visual and olfactory cues have been effectively used to monitor oriental fruit flies infestation (Jiang et al., 2024) [48] . Rajashekhar et al., 2024 [83] demonstrated that pheromone-based monitoring integrated with biological, botanical and need-based insecticide applications effectively managed Spodoptera frugiperda S. in maize. The IPM strategy increased yield by 17.49-24.87 percent over farmers' practices.

Recent strategies in Semiohemicals Application in Integrated Pest Management 1) Push and pull strategyThe term push-pull was first conceived by Pyke et al., (1987) [81] as a strategy for insect pest management. Push-pull experiments using a combination of repellents and attractants could be considered a sustainable alternative to conventional insecticides. Repellents could 'push' away insects from vulnerable crops whereas at the same time attractants could 'pull' them into annihilating traps (Cook et [27,54,86,94] . Push-pull and habitat management are core components of Integrated Pest Management (IPM) programs that aim to synergize multiple compatible tactics for long-term pest control (Barzman et al., 2015) [14] . Pull plants attract insect pests using volatile compounds, visual signals, and arrestant cues. They may also release oviposition stimulants and offer better nutrition to maintain the pests (Eigenbrode et al., 2016) [34] . In the maize-growing farming systems of East Africa, combining push-pull with peripheral vegetation like hedgerows and woodlots has enhanced the benefits of pest and weed control, while offering additional ecosystem services such as soil and water conservation, fodder production, and carbon sequestration (Khan et al., 2016) [55] . Use of the push-pull semiochemical technique is effective in reducing Ips (-66 percent) and Dendroctonus (-54 percent) compared to control;

(2) Among push-pull treatment sources, conspecific semiochemicals combined with heterospecific and NHVs provide the maximum reduction for Dendroctonus (Afzal et al., 2023) [6] . [53] Maize Spodoptera frugiperda Desmodium spp.

Napier grass / Brachiaria spp.

Reduced fall armyworm infestation and plant damage; increased maize yield Midega et al. (2018) [65] Maize Stemborers Desmodium spp.

Napier grass Repulsion of adult moths from maize and increased mortality of larvae on trap plants Khan et al. (2008) [52] Cotton

Helicoverpa armigera

Neem-based repellent Pigeon pea / okra Repels oviposition from cotton and concentrates pest activity on trap crops Duraimurugan and Regupathy (2005) [33] Likewise, combining push-pull with flower strips and beetle banks has increased the abundance and diversity of natural enemies in brassica crops in New Zealand, resulting in improved biological control of aphids and moths (Tompkins, 2010) [104] .

2) Lure and infect

In recent years, the use of semiochemicals combined with entomopathogenic fungi has emerged as a promising strategy for pest management in agriculture. This method is often referred to as "lure and infect" and it utilizes natural dispersal mechanisms of pathogens while increasing their effectiveness through targeted application (Vega et al., 2009) [107] . The lure-and-infect (also known as attract-andinfect) strategy has demonstrated promising capacity for controlling several dipteran pests, including tsetse flies and Mediterranean fruit flies (Maniania, 2002 [31,61,70] . Applying this strategy to Beauveria bassiana B. presents a major technical challenge: many highly potent attractant components or their associated chemical environments can suppress conidial germination and mycelial growth of B. bassiana (Toledo et al., 2007; Khun et al., 2021) [56,103] . A study by Mfuti et al., (2017) [64] investigated the combined use of Metarhizium anisopliae M. and the thrips attractant Lurem-TR for managing bean flower thrips on cowpeas. Their field experiments compared spot and cover spray applications, where the spot application ("lure-and-infect") was the most effective. Thrips density was reduced from 67.4 ± 10.30 thrips per plant in the control to 30.5 ± 6.0 thrips per plant. Importantly, this method required 81.2 per cent less inoculum than cover sprays, making it a cost-effective approach for small-scale farmers. Dembilio et al., (2018) [29] developed a lure and-infect device using the red palm weevil aggregation pheromone to attract adults and Beauveria bassiana to infect them. Laboratory, semi-field and field studies demonstrated the potential of the device for controlling the red palm weevil. Metarhizium brunneumbased lure-and-infect device effectively suppressed Drosophila suzukii M. The treatment caused 48percent mortality in untreated males and reduced female fecundity by 84.7 percent, indicating its potential as an eco-friendly IPM strategy (Yousef et al., 2018) [118] .

3) Integration of semiochemicals and remote sensing technologies

Researchers have employed drones equipped with sensors to detect volatile organic compounds (VOCs) emitted by plants under stress. These VOCs serve as semiochemicals that can indicate factors such as drought, nutrient deficiencies, or pest damage. By combining this technology with remote sensing data on vegetation health, researchers can assess the physiological condition of crops or natural vegetation with high spatial resolution (Lasa et al., 2024) [58] . Bapatla et al., (2026) [12] integrated pheromone-trap monitoring with Landsat-8 remote sensing to monitor the yellow stem borer, Scirpophaga incertulas, in rice fields of Srikakulam, Andhra Pradesh. The study used pheromone traps along with NDVI and land surface temperature (LST) to identify pest hotspots. Smart trap containing a pheromone-kairomone lure and automatic imaging for remote monitoring of codling moth in apple and pear orchards. The system allowed trap catches to be monitored remotely, reducing the need for frequent manual inspection (Preti et al., 2021) [79] .

Conclusion

Semiochemicals are important behaviour-based tools in IPM that manipulate insect behaviour for detection, monitoring and pest suppression. Pheromones and allelochemicals are used in mass trapping, mating disruption, push-pull strategies, host-marking manipulation and lure-and-infect systems. Successful examples in brinjal, cereals and lepidopteran pests show that they can effectively reduce pest populations when combined with other IPM methods. Recent advances such as controlled-release formulations, automated traps, IoT monitoring and remote sensing are making semiochemical-based management more precise, scalable and timely. However, challenges remain regarding formulation stability, release rate, trap design, environmental variation, cost and compatibility with biological control agents. Overall, the future of semiochemical-based IPM depends on integrating chemical ecology, biological control, digital technologies, remote sensing and precision agriculture. This integration can provide targeted, economical, environmentally safe and climate-resilient pest management, provided that technologies are further validated under field conditions and made accessible to farmers

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