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",isbn:null,printIsbn:null,pdfIsbn:null,doi:null,price:0,priceEur:null,priceUsd:null,slug:null,numberOfPages:0,isOpenForSubmission:!1,hash:"cc796459268324e827219d1d904e4265",bookSignature:"Prof. Moulay Tahar Lamchich",publishedDate:null,coverURL:"https://cdn.intechopen.com/books/images_new/7196.jpg",keywords:"Induction motor, smart motor, electrical vehicles, energy generation, drives, electromechanical, hybrid transportation, smart control, high efficiency motor, variable speed drives, power electronic, energy efficiency.",numberOfDownloads:null,numberOfWosCitations:0,numberOfCrossrefCitations:0,numberOfDimensionsCitations:0,numberOfTotalCitations:0,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"July 4th 2018",dateEndSecondStepPublish:"July 25th 2018",dateEndThirdStepPublish:"September 23rd 2018",dateEndFourthStepPublish:"December 12th 2018",dateEndFifthStepPublish:"February 10th 2019",remainingDaysToSecondStep:"3 years",secondStepPassed:!0,currentStepOfPublishingProcess:5,editedByType:null,kuFlag:!1,biosketch:null,coeditorOneBiosketch:null,coeditorTwoBiosketch:null,coeditorThreeBiosketch:null,coeditorFourBiosketch:null,coeditorFiveBiosketch:null,editors:[{id:"21932",title:"Prof.",name:"Moulay Tahar",middleName:null,surname:"Lamchich",slug:"moulay-tahar-lamchich",fullName:"Moulay Tahar Lamchich",profilePictureURL:"https://mts.intechopen.com/storage/users/21932/images/system/21932.png",biography:"Moulay Tahar Lamchich is a Professor at the Faculty of Sciences Semlalia at Marrakech (Morocco). He completed his thesis in electromechanics in September 1991 and received his third cycle degree. Dr. Lamchich received his Ph.D. from the same university in July 2001. His main activity is based on short-circuit mechanical effects in substation structures, control of different types of machine drives, static converters, active power filters. In the last decennia, his research interests have included renewable energies, particularly the control and supervision of hybrid and multiple source systems for decentralized energy production, and intelligent management of energy. He has published more than fifty technical papers in reviews and international conferences. With IntechOpen, he has published two chapters and was editor of the books “Torque Control” and “Harmonic Analysis”. 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From chapter submission and review, to approval and revision, copyediting and design, until final publication, I work closely with authors and editors to ensure a simple and easy publishing process. I maintain constant and effective communication with authors, editors and reviewers, which allows for a level of personal support that enables contributors to fully commit and concentrate on the chapters they are writing, editing, or reviewing. I assist authors in the preparation of their full chapter submissions and track important deadlines and ensure they are met. I help to coordinate internal processes such as linguistic review, and monitor the technical aspects of the process. As an ASM I am also involved in the acquisition of editors. 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Venkateswarlu",coverURL:"https://cdn.intechopen.com/books/images_new/371.jpg",editedByType:"Edited by",editors:[{id:"58592",title:"Dr.",name:"Arun",surname:"Shanker",slug:"arun-shanker",fullName:"Arun Shanker"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"72",title:"Ionic Liquids",subtitle:"Theory, Properties, New Approaches",isOpenForSubmission:!1,hash:"d94ffa3cfa10505e3b1d676d46fcd3f5",slug:"ionic-liquids-theory-properties-new-approaches",bookSignature:"Alexander Kokorin",coverURL:"https://cdn.intechopen.com/books/images_new/72.jpg",editedByType:"Edited by",editors:[{id:"19816",title:"Prof.",name:"Alexander",surname:"Kokorin",slug:"alexander-kokorin",fullName:"Alexander Kokorin"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"314",title:"Regenerative Medicine and Tissue Engineering",subtitle:"Cells and Biomaterials",isOpenForSubmission:!1,hash:"bb67e80e480c86bb8315458012d65686",slug:"regenerative-medicine-and-tissue-engineering-cells-and-biomaterials",bookSignature:"Daniel Eberli",coverURL:"https://cdn.intechopen.com/books/images_new/314.jpg",editedByType:"Edited by",editors:[{id:"6495",title:"Dr.",name:"Daniel",surname:"Eberli",slug:"daniel-eberli",fullName:"Daniel Eberli"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"57",title:"Physics and Applications of Graphene",subtitle:"Experiments",isOpenForSubmission:!1,hash:"0e6622a71cf4f02f45bfdd5691e1189a",slug:"physics-and-applications-of-graphene-experiments",bookSignature:"Sergey Mikhailov",coverURL:"https://cdn.intechopen.com/books/images_new/57.jpg",editedByType:"Edited by",editors:[{id:"16042",title:"Dr.",name:"Sergey",surname:"Mikhailov",slug:"sergey-mikhailov",fullName:"Sergey Mikhailov"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"1373",title:"Ionic Liquids",subtitle:"Applications and Perspectives",isOpenForSubmission:!1,hash:"5e9ae5ae9167cde4b344e499a792c41c",slug:"ionic-liquids-applications-and-perspectives",bookSignature:"Alexander Kokorin",coverURL:"https://cdn.intechopen.com/books/images_new/1373.jpg",editedByType:"Edited by",editors:[{id:"19816",title:"Prof.",name:"Alexander",surname:"Kokorin",slug:"alexander-kokorin",fullName:"Alexander Kokorin"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}}]},chapter:{item:{type:"chapter",id:"68636",title:"Potential Roles of Matrix Metalloproteinases in Malignant Mesothelioma",doi:"10.5772/intechopen.88783",slug:"potential-roles-of-matrix-metalloproteinases-in-malignant-mesothelioma",body:'Malignant mesothelioma (MM) is a rare, aggressive cancer that originates from mesothelial tissue in the pleura, peritoneum, and pericardium; MM has been associated with asbestos exposure, especially in occupational settings [1]. In some countries, such as Turkey and Japan, MM is also due to environmental asbestos exposure, which affects people who live in the vicinity of natural asbestos mines or factories that use asbestos [2, 3, 4]. Mesothelioma is highly resistant to conventional cancer therapies. MM patients usually have a poor prognosis, with a median survival of 12–18 months, due to the lack of effective treatments and difficulty in diagnosing this disease at the early stage [5, 6, 7]. In general, there are three main histological subtypes of mesothelioma. The epithelioid and sarcomatoid subtypes are characterized by cuboid and fibroblastoid cells, respectively. The biphasic subtype contains a mixture of both cell types and confers the worst prognosis. The most widely used treatments for MM are surgery with or without adjuvant chemotherapy and/or radiotherapy [8]. The first-line treatment option for unresectable MM is chemotherapy with cisplatin plus pemetrexed [9, 10]. Nevertheless, MM may be resistant to these conventional therapeutic approaches, and palliative care strategies are controversial. Although crocidolite and/or chrysotile have not been used for more than 10 years in many developed and developing countries, high mortality rates associated with mesothelioma persist since the clinical manifestations of MM are insidious and nonspecific. It is worth noting that MM has a long latency period (mean, 30–40 years) from the time of asbestos exposure to tumorigenesis [4, 11]. Thus, valuable biomarkers for the prediction or diagnosis of MM at early stages, prognostic markers, and novel therapeutic strategies are urgently needed.
Matrix metalloproteinases (MMPs, also known as matrixins) are a family of zinc-dependent endopeptidases that degrade all components of the extracellular matrix (ECM); thus, MMPs are involved in ECM remodeling. In addition to functioning as the main ECM regulators, MMPs also modulate intra- and extracellular signaling pathways and networks through the proteolytic processing of various biomolecules. The first MMP was reported by Gross and Lapiere as a collagenase engaged in tail resorption during tadpole metamorphosis [12]. To date, 24 MMP genes, including a gene duplication, that encode 23 unique MMP proteins have been identified in humans [13, 14]. According to substrate specificity, sequence similarity, and specific role, MMPs can be divided into eight main groups: (1) collagenases (MMP-1, MMP-8, and MMP-13), (2) matrilysins (MMP-7 and MMP-26), (3) metalloelastase (MMP-12), (4) stromelysins (MMP-3, MMP-10, and MMP-11), (5) gelatinases (MMP-2 and MMP-9); (6) enamelysin (MMP-20); (7) membrane-type MMPs (MMP-14, MMP-15, MMP-16, MMP-17, MMP-24, and MMP-25), and (8) others (MMP-19, MMP-21, MMP-23, MMP-27, and MMP-28) [13, 15]. Interestingly, the proteolytic activities of MMPs are precisely controlled by activation of their precursors and inhibition by endogenous inhibitors, α-macroglobulins, and tissue inhibitors of metalloproteinases [13]. Except for six membrane-associated MMPs, the other 17 MMPs are soluble secreted enzymes [16]. In addition, growth factors, chemokines, and cytokines modulate the expression of MMPs through various pathways to affect ECM degradation and, in turn, influence growth factors, which ultimately affect cancer cell migration and invasion [17, 18].
Of note, MMPs are expressed in various cancer tissues, and their expression levels are closely associated with the properties of invasive growth and metastasis [15]. Accumulating evidence suggests that ECM degradation by MMPs at the cell surface enhances tumor growth, invasion, and metastasis through the proteolytic degradation of ECM, altered cell-cell, and cell-ECM interactions and effects on cell migration and angiogenesis [17, 19]. More recently, the roles of different MMPs have become increasingly studied in the field of MM research. Experimental evidence indicates that MMP-1, MMP-2, and MMP-9 are involved in mesothelial carcinogenesis. Several MMPs, such as MMP-7, MMP-14, and MMP-9, are potential biomarkers for MM. In the following sections, we will describe the roles of MMPs in carcinogenic mechanisms based on in vivo and in vitro experimental evidence, outline the clinical findings, and highlight the possible roles of MMPs in MM, as well as future prospects.
Some MMPs are upregulated and considered mesenchymal markers of epithelial-to-mesenchymal transition (EMT), such as MMP-1, MMP-2, and MMP-9 [20]. EMT not only is associated with many physiological processes, such as embryonic development, but also plays a vital role in pathological processes, including cancer cell invasion and migration [21, 22, 23]. During EMT, epithelial cells lose their phenotype and acquire a mesenchymal phenotype, including the loss of cell polarity and cell adhesion in cell-cell and cell-basement membrane interactions and the acquisition of ECM degradation ability, which is directly related to MMPs. Currently, published studies implicate MMPs as inducers of EMT during MM progression. In addition, MMPs play a mediator role in cellular signaling pathways controlled by growth factors and cytokines [17, 24]. Here, we describe these two main roles of MMPs in MM carcinogenesis. Moreover, we propose possible mechanisms involving MMPs, as shown in Figure 1.
Schematic representation of MMP-involved mechanisms in MM carcinogenesis. See detail in text. MM, malignant mesothelioma; EMT, epithelial-to-mesenchymal transition; ECM, extracellular matrix.
MMP-1 is an interstitial collagenase that specifically targets the degradation of collagen types I–III [25]. Schelch et al. reported that in malignant pleural mesothelioma (MPM) cells in vitro, fibroblast growth factor 2(FGF2) and epidermal growth factor (EGF) may induce EMT via mitogen-activated protein kinase kinase (MEK)/MMP-1 signaling [26]. The experimental results indicated that MMP-1 inhibition by the pan-MMP inhibitor GM6001 or transfection with siRNAs targeting MMP-1 could prevent FGF2-induced cell scattering and invasion in the M38K cell line (a biphasic MPM cell line) [26]. In MPM tissue specimens, higher MMP-1 expression was observed in the sarcomatoid compartment than in the epithelioid compartment. Normal pleura were weakly positive for MMP-1 [26]. These results suggest that MMP-1 causally contributes to sarcomatoid morphology and increases cell invasiveness during EMT.
MMP-2, also named gelatinase A, is expressed by almost all cell types, and its classical substrates are denatured collagen (gelatin) and basement membrane [25, 27]. Indeed, MMP-2 acts as a cancer-associated EMT inducer or modulator in a number of tumors, such as breast cancer [16, 28], hepatocellular carcinoma [29], prostate cancer [30], ovarian cancer [31], oral squamous cell carcinoma [32], and MM [33]. Regarding MM, MMP-2 secretion from human normal mesothelial MeT-5A cells increased upon treatment with chrysotile or transforming growth factor-β (TGF-β) [33], and EMT was induced. This in vitro experimental result of increased MMP-2 secretion by cells exposed to chrysotile asbestos suggests changes in the surrounding microenvironment that render the ECM more amenable to degradation and invasion [33, 34]. Of course, the underlying mechanism of MMP-2-induced EMT in MM development requires further study.
MMP-9 is a type IV collagenase also known as gelatinase B [35] that has a similar ability to cleave gelatin as MMP-2. MMP-9 has been recognized as an EMT mediator in cancer progression and appears to be a potential therapeutic target [35, 36, 37]. Elevated MMP-9 levels were observed in a 3D microtumor model of patient-derived mesothelioma cells, consistent with the elevated MMP-9 levels in patient breast tumors compared to healthy mammary glands [38]. Moreover, MMP-9 secreted into conditioned media by large microtumors induced a migratory phenotype in nonmigratory small microtumors, and blocking MMP-9 with GM6001 effectively abolished the collective migration of mesothelioma microtumors [38]. These findings imply that a self-regulated positive feedback loop involving MMP-9 is established during tumor progression and migration [38]. Additionally, the invasion of H2052 (mesothelioma cell line) and JP5 cells (primary mesothelioma cell line) into a 3D collagen matrix induced by gremlin-1 (a protein antagonist of bone morphogenetic proteins) was significantly alleviated by GM6001 and BB2516 (broad-spectrum MMP inhibitors) [39]. Interestingly, in our previous study, we found that serum MMP-2 and MMP-9 levels were correlated with each other in both healthy control and MM groups in a Han cohort from Eastern China [40]. Nevertheless, there were no significant differences in MMP-2/MMP-9 levels between the healthy control and MM groups.
Various growth factors, cytokines, and miRNAs engage specific cellular signaling pathways, such as the MEK and extracellular signal-regulated kinase (ERK) signaling pathways, to regulate MMP expression levels to degrade the ECM, and MMPs then contribute to the release of tumor-related factors, such as vascular endothelial growth factor and TGF-β, from the ECM [17, 24].
MMP-1 expression showed an increasing trend in MM cell lines from no treatment to treatment with FGF2 and EGF and a pronounced decrease upon treatment with selumetinib (MEK inhibitor), suggesting that the growth factors FGF2 and EGF regulate MMP-1 expression via the MEK signaling pathway in MM [26]. TGF-β, another important growth factor that regulates cell growth and differentiation, affects MMP-2 expression in MeT-5A [33] and JL-1 cells [39]. Moreover, growth hormone-releasing hormone (GHRH) antagonists (MIA-602 and MIA-690) equally blunted MMP-2 and MMP-9 mRNA levels in both REN and MSTO-211H cells (MM cell lines), indirectly indicating that MMP-2/MMP-9 expression is induced by GHRH [41], as well as by adenosine diphosphate in ZL55 cells (an epithelioid MM cell line), via the nuclear factor kappa-B, protein kinase B, and ERK1/2 signaling pathways [42]. Interestingly, microtumor treated with GM6001 showed reduced pERK/ERK ratios and ERK activation [38]. Notably, miR-591 targets MMP-2 expression, and overexpression of miR-591 inhibited MMP-2 levels in MPM cells [43]. These experimental results show that MMP expression is regulated by various factors via multiple signaling pathways and that MMPs interact with such inducers and signaling pathways in MM carcinogenesis.
To date, MM is still difficult to diagnose in early stages due to our limited knowledge of its molecular pathogenesis. Indeed, pathological examination techniques to diagnose MM and distinguish MM from other diseases must be improved [44]. However, more molecular markers are required to distinguish benign from malignant mesothelial disease or other tumors. In addition, effective pathologic predictors of prognosis and therapeutic response are urgently needed. Since MMPs are involved in tumor pathogenesis, some MMPs may be potential pathological markers.
In general, MMP expression and activation are very low and tightly regulated during normal tissue homeostasis. MMP production and activation are rapidly induced during active tissue remodeling and in pathological conditions such as cancer [37]. MMP-7 and MMP-14 are potential diagnostic and prognostic biomarkers of mesothelioma, respectively. MMP-7 is a highly specific negative biomarker to distinguish MM from other high-grade serous carcinomas with 100% specificity and moderate sensitivity, but it cannot distinguish mesothelial cells from reactive mesothelial cells in serous effusion due to uniformly negative expression of MMP-7 in reactive mesothelial cells [45]. It is intriguing that MMP-14 is a potential biomarker for the differential diagnosis of MPM and reactive mesothelial hyperplasia (MH). A group from Italy found that MMP-14 expression is markedly increased in MPM patient specimens compared with MH specimens based on polymerase chain reaction array and immunohistochemistry analyses [46]. MMP-14 levels have been reported to be elevated in all tissue samples from MM patients compared to those from normal individuals, but more evidence is needed to substantiate MMP-14 as a diagnostic biomarker for MM [47]. MMP-14 expression has prognostic value for MM. Clinically high MMP-14 expression in MM patients is significantly correlated with poor prognosis [47].
Although most mesotheliomas are attributable to asbestos exposure, genetic factors are also important causes of carcinogenesis. Gene mutations influence the prognosis of MM. For example, heritable mutations in BRCA1-associated protein-1 (BAP1), a tumor suppressor gene, may predispose individuals to asbestos-related MM [48, 49]. Moreover, Baumann et al. reported that mesothelioma patients with germline BAP1 mutations have a seven-fold improvement in long-term survival [50].
More recently, some MMP single-nucleotide polymorphisms (SNPs) have been found to have potential as genetic biomarkers for MM. For instance, Štrbac et al. reported that patients carrying a polymorphic MMP-9 rs2250889 allele had a negative outcome, with a shorter time to progression (TTP) (6.07 vs. 10.03 months, HR = 2.45, 95% CI = 1.45–4.14, p = 0.001) and worse overall survival (OS) (9.23 vs. 19.2 months, HR = 2.39, 95% CI = 1.37–4.18, p = 0.002) than those with the reference allele [51]. However, patients harboring at least one polymorphic MMP-9 rs20544 allele had a positive outcome, with a longer TTP (10.93 vs. 9.40 months, HR = 0.57, 95% CI = 0.38–0.86, p = 0.007) and improved OS (20.67 vs. 13.50 months, HR = 0.56, 95% CI = 0.37–0.85, p = 0.007) [51]. These researchers also found that the MMP-2 rs243865 polymorphism plays a protective role in MM; carriers of this polymorphism have a decreased risk for MM (OR = 0.66, 95% CI = 0.44–1.00, p = 0.050) [52]. Interestingly, the decreased risk for MM is more pronounced in people exposed to asbestos [52]. These findings provide insight into some MMP SNPs that are considered genetic biomarkers, indicate the prognosis of MM patients, and predict susceptibility to MM. In the future, appropriate genetic counseling and clinical management should be considered for MM patients who are carriers of MMP-2/MMP-9 susceptibility SNPs.
In this chapter, we provide an overview of recent findings on MMP function in MM and the mechanisms by which MMPs may induce both phenotypic and genotypic alterations that facilitate MM progression and invasion. Accumulating evidence indicates that tumor-associated MMPs can stimulate processes associated with EMT, a developmental event that is activated in MM cells during invasion and metastasis. Meanwhile, future investigations on extracellular targets and intracellular signaling pathways through which MMPs can induce EMT of MM cells will provide insight into novel therapeutic targets. We also describe possible roles of MMPs as pathological markers or genetic biomarkers in MM. Certainly, the underlying mechanisms of secreted MMPs, including their function and circulation, are complex in MM and remain to be elucidated in the future.
This study was supported by the National Natural Science Foundation of China (81502794, 81973011), the Qianjiang Talents Project of Zhejiang Province (QJD1602027), the Science and Technology Department of Zhejiang Province (1925D), the Zhejiang Provincial Natural Science Foundation of China (LY16H240001), the Health Commission of Zhejiang Province (2017PY015, 2018KY038), and the Science and Technology Project of Cixi City (CN2018023). The research assistance of Dr. Xing Zhang and Dr. Jianlin Lou is gratefully acknowledged.
The authors declare no conflicts of interest.
Many insects are called flies such as butterflies and dragonflies, but only insects belonging to the order diptera are known as “true flies.” Dipteran insects (flies and mosquitoes) are holometabolous insects which means they have complete metamorphosis life cycle. Dipteran insects have 4 stages in their life cycle (adult, pupae, larvae and egg). Name of larval stages of these insects is “maggot”. Adults of this order are recognized according to wing types. Front wings are developed for flying. Other pairs of wings are undeveloped and they have balancing function when insects fly. These type wings give order its name: two (di-), wings (ptera). One pair of wing provides flying of insects and other pair developed into balancing stuructures. The name of other pair of wings which provide balancing during flight, called “halteres”. Except mosquitoes, dipteran insects have sponging-sucking mouthparts. Mosquitoes and some other have piercing-sucking mouthparts. The Diptera order is divided into two or three subclasses: Nematocera and Brachycera, the second being Orthorrhapha and Cyclorrhapha [1]. There are approximately 152.000 identified species in this order which are distributed about 130 families [2]. Houseflies, hoverflies, mosquitoes and fruit flies are the most important species which belong to the diptera order. Houseflies carry diseases such as cholera and dysentery. The fly cause serious problems by carrying disease organisms onto food. They take disease organisms from leg hair or eat them and then feed them to other foods. The adults of hoverflies feed mainly on pollen grains. The larvae of some species eat rotten plant and animal materials in the soil or in ponds, lakes and streams.
\nAnother species of hoverflies’s larvae are predators and feed on some harmful insects such as aphids and thrips etc. So, the larvae of these species are important for biological control as natural enemies.
\nThe females of mosquitoes suck the blood of human and several animals such as farm animals and wild animals. Mosquitoes are vector insects and cause the spread of serious illnesses such as malaria and leishmaniasis etc. [3]. Fruit flies are the most destructive pests in horticultural growing. For example, Olive fruit fly are the main pest in growing areas where table and olive oil production are made throughout the world.
\nGeneral objectives of this chapter is to provide information about the habitats, feeding patterns and life cycles of insects belonging to the order Diptera, and the stages in which they exist in their life cycle; However, it is to provide information on the characteristics of each stage (adult, egg, larva and pupa) in the life cycle of these insects. In addition, to give information about the role of insects in this order for human and animal health and agricultural production.
\nDipteran order has 125.000 insect species and is one of the biggest order throughout the world and is highly diverse. Our world score is based on data from higher than 152.000 known species and higher than 130 identified families and these data are from the“World Diptera’s Bio Systematic Database” [2]. Diptera order has the fourth place after the Lepidoptera, Hymenoptera and Coleoptera. Many species belonging to the order Diptera are found in almost all zoogeographic regions of the world. These species are well adapted to a broad range of habitats. Except the depth of oceans, they can live many habitats on earth [1]. Maximum nutrient and biomass formation occurs in the maggots of Dipteran order, and the adult this order usually receives the energy which they need to feed their muscles which have functions for flying. Widely the broad range of insects (flies) looking for food, the food of those insects includes honey extract or nectar, blood of vertebrate, pollen, hemolymph of other insects, and another liquefied or liquefied biological resources which are suspended or dissolved in vomiting fluid or saliva. Few groups of adults are predators. Other few groups are completely devoid of mouthparts, so they do not receive food. Therefore, they have short life span. Larval stage of many species, as they exist, live in water, alive plant tissues and rotten organic matter. Moreover, they live as parasites or parasitoids of several animals. These larvae need a humid and wet atmosphere. In addition, the eggs these species hatch in water surfaces and larval stage occurs on the water surfaces. Maximum larvae live freely, also can be found in water, sediments, trees, fruits or decaying biological material, while other larvae are found in the tissues of living beings [2].
\nDiptera insects are holometabolous (complete metamorphosis) which means that they have 4 stages in their life cycles (adult, egg, larvae and pupae). Adult females lay their eggs and the amounts of eggs vary according to species. The females of some species lay a few eggs, some other lay thousands of them. Generally, females lay their eggs near the water and lay them as a group or singly.
\nThe eggs are laid grouped or individually by adult females, and the females usually lay their eggs in water and are sometimes attached to materials. Except for diapause eggs, eggs tend to stand only last for a few days, which are used to prevent lack of water or unwanted temperatures in the ecosystem [4].
\nMosquitoes which belong to the order of diptera and Culicidae family, lay their eggs in bunches or can lay singly in or near water (Figure 1). In contrast, some dipteran insects such as Olive fruit fly (
Standing water mosquito and eggs.
Egg of olive fruit fly inside olive fruit.
Usually, adult flies lay their eggs, which will pass into the larval stage within a few hours or days after hatching. Amounts of eggs which are laid by adult females vary between 1 and 250.However, multiple sequential egg batches can be made. Females of Medfly lay 300 eggs in her lifetime. Besides, the Green bottle fly (
Larvae of dipteran insects are easily known by the absence of thoracic structures (legs). This type of larvae is called Apod larvae. Body parts of larvae are usually fleshy (thorax and abdomen). The whole body of larvae is tubular and long. Larvae are approximately 2 0r 25 mm, but some species can be reached about 10 cm length. There are 12 segments in the bodies of larvae and that segmentation pattern is most common. 3 of them is found in thorax and 9 of them in the abdomen [4]. Some true midges, commonly seen in anoxic habitats such as blood worms, have an pigment which has invertebrate form and respiration function. This is called “hemoglobin” that helps to capture oxygen molecules [4] (Figure 3). After the eggs hatch, the larvae of the maximum species pass 3 to 4 stages on land or close to the bottom or above the water surface. For instance, females of Olive fruit fly And Spotted wing Drosophila lay their eggs inside the developing fruit [5] (Figures 4 and 5). After the emerging of larvae, they consume the rotten material in which they are laid. They eat more foods to store energy and nutrients for pupa stage [1]. Larval period is completed from nearly 2 weeks to several months. Larvae of Diptera insects do not have wing pads but are found in pupae [4]. Respiration process takes place above the skin of many larvae of dipteran insects. There are small gills above the skin of some taxa. Other larvae of dipteran insects have spikes and they absorb oxygen from atmosphere by using long or short breathing tubes.
\nPlaces of respiratory spikes in Diptera larva. (A) Peripneustic, (B) Amphipneustic, (C) Metapneustic, (D) Apneustic.
Larva of the olive fruit fly.
Spotted wing drosophila: (A) egg-laying areas and (B) larvae of spotted wing drosophila.
Pupae of dipteran insects have a non-functional mandible (adecticeous). Their appendages can be independent from their body (externally) or attached to the body (obtect). Pupae of exarate types are hidden inside the hardened skin (puparium) of the final stage of larvae [2].
\nThe pupae stage of dipteran insect varies significantly in shape. The pupae of some flies look like cross shape between larvae and adult, while other pupae shape of flies are featureless and they have a structure similar to seeds. The first forms are typical for Nematocera and are defined as having obtect or body-attached appendages. For example, the pupae of a Crane fly (Tipulidae) have identifiable head, thorax and abdomen, but covers of antennae, legs and wing pads attach to the body of pupae. The exterior of the Nematocera pupa can be decorated with spines, breathing apparatus which are similar to gill or locomotory paddles.
\nBrachycera and Cyclorrhapha form the pupal stage in a different and more discreet way. The so-called higher Diptera family produces pupae that are described as coarctates, meaning “compressed” or “constricted”. These taxa (eg Syrphidae, Drosophilidae, Muscidae) form a puparium consisting of hardened skin of the late larval stage [6] (Figure 6). The pupation of some flies occurs in the olive fruit or under the soil [5]. After the oviposition, the eggs hatch and the larvae feed on the fleshy part of the fruit, but leaves the fruit when ripe and some continue to appear inside the fruit. Larvae fall to the ground and pupation takes place in the soil [7].
\nFemale, serrate ovipositor, male with typical black spots, larvae inside the fruit and pupae of
Adults of dipteran insects have segmented body which includes head, thorax and abdomen parts. They have compound eyes which are found both side of head [1]. The size and shape of compound eyes are highly variable.
\nAdults have dark reddish to black color compound eyes. Some families of dipteran insects have crossbands or spots of different colors such as Tabanidae, Syrphidae, Tephritidae, Sciomyzidae families [8].
\nThere is small space in front of the head of adult dipterans. The function of this small space is to help the adults to see wider area when insects fly. Body color of adults changes from brown to black, orange or yellow, depending on the dipteran species [1]. For instance, body color of adults of
General view of fruit fly adults.
Thorax color of Spotted wing Drosophila is pale brown and there are horizontal black lines in the abdomen. Males have spots on their wings. Adult males of the
Spotted wing drosophila: (A) the black spots in the wings of male, (B) female, (C) egg laying organ (ovipositor) of female.
Length range varies of adults varies according to dipteran species between 1 to 12 mm, but relative huge species are between 25–60 mm [4]. Sensory organ is found in front of the head of adults which is called antennae. Antennae are filiform and front wings are developed to fly and hind wings are undeveloped (halteres) to balance when insects fly and tarsi 5-segmented [2].
\nDipteran order has 125.000 insect species and is one of the biggest order throughout the world and is highly diverse. Diptera order has the fourth place after the Lepidoptera, Hymenoptera and Coleoptera. Houseflies, hoverflies, mosquitoes and fruit flies are the most important species which belong to the diptera order. Fruit flies cause destructive damages in agricultural production. Besides, houseflies transmit serious diseases transmits diseases by carrying disease organisms onto food such as cholera and dysentery. Mainly, adult hoverflies consume pollen grains. Larvae of some species feed on rotten plant and animal materials in lakes, streams and ponds or inside the soil. Another species of hoverflies’s larvae are predators and feed on some harmful insects such as aphids and thrips etc. So, the larvae of these species are used in the biological control of harmful insects as natural enemies. The females of mosquitoes have piercing-sucking mouthparts and suck the blood of human, livestock and animals by using their mouthparts. Mosquitoes are vector insects and cause the spread of serious illnesses such as malaria and leishmaniasis etc. Many of the flies which belong to Tephritidae and Drosophilidae families, cause serious damages in the agricultural production such as Olive fruit fly, Medfly, Peach fruit fly, Celery fly Spotted wing drosophila. Dipteran insects can be recognized by some features such as developed membranous front wings and hind wings are undeveloped and called “halteres” which have functions as balancing when insects fly. Dipteran insects have complete metamorphosis (holometabolous) life cycle which means that there are 4 stages (egg, larvae, pupae, adult). Females of the adults lay their eggs into the food source or water. Eggs hatch and larvae complete their development. Pupation occurs under the soil, plant and animal tissues and water. The richness of the species in this order, the living in different ecological conditions and the morphological differences show people that this order is economically important and externalizes the diversity of invertebrate creatures in the world. There are insects in the Diptera order that cause serious problems in human, animal health and agricultural production. It is important to know the life cycles and habitats of these insects. Accordingly, the issues are followed (life cycle and habitats) to minimize disease transmission and damage in agricultural production. Another studies which will do in the future, will be useful in determining the economic effects of diptera insects on human health and agricultural production.
\nAuthor has no conflict of interest.
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